Algal species composition and phosphorus recycling at contrasting grazing pressure: an experimental study in sub-Antarctic lakes with two trophic levels

1. To test the response of algal communities to altered grazer abundance in lakes lacking efficient predators on herbivores, we performed field and experimental studies in two sub-Antarctic lakes (South Georgia).
2. The number of algal species in these high latitude lakes is low, and all dominant species have grazer-resistant adaptations, including spines in three dimensions ( Staurastrum sp.), large size ( Tribonema sp.), a mucus sheet allowing viable passage through the gut ( Chlamydocapsa sp., Elakatothrix sp.) or ability to recruit individuals from the sediment surface ( Mallomonas sp.).
3. Algal community composition was only slightly changed by experimentally altered grazer abundance, indicating that it was already adapted for a high grazing pressure. Hence, the diets of herbivores were restricted to vulnerable food organisms such as Mallomonas sp. and heterotrophic flagellates in the water column, and to benthic food sources.
4. At high grazer abundance, the concentration of available phosphorus (PO₄-P) in the water was lower than at low grazer abundances, due to inefficient nutrient regeneration by the copepod herbivores. Hence, in lakes where copepods are dominant grazers, algae suffer both directly from grazing and indirectly from reduced nutrient availability.     

Quantification of invertebrate predation and herbivory in food chains of low complexity

Zooplankton grazing impact on algae, heterotrophic flagellates and bacteria, as well as invertebrate predation on herbivorous zooplankton, were investigated in two sub-Antarctic lakes with extremely simple food chains. The two species of herbivorous zooplankton present in the lakes (the copepods boeckella michaelseni and Pseudoboeckella poppei) exerted substantial grazing pressure on algae. However, the dominant algal species exhibited properties that enabled them to avoid (large size or extruding spines, e.g. Staurastrum sp., Tribonema sp.) or compensate (recruitment from the sediment, Mallomonas sp.) grazing. There are only two potential invertebrate predators on the herbivorous copepods in the two lakes: the copepod Parabroteas sarsi and the diving beetle Lancetes claussi. Vertebrate predators are entirely abscent from sub-Antarctic lakes. Based on our experiments, we estimated that the predators would remove at most about 0.4% of the herbivorous copepods per day, whereas planktivorous fish, if present in the lakes, would have removed 5–17% of the zooplankton each day. Consequently, the invertebrate predators in these high-latitude lakes had only a marginal predation impact compared to the predation pressure on zooplankton in the presence of vertebrate predators in temperate lakes. The study of these simple systems with only two quantitatively functionally important trophic links, suggests that high grazing pressure foreces the algal community towards forms with grazer resistant adaptations such as large size, recruitment from another habitat, and grazer avoidance spines. We propose that due to such adaptations, predictions from food web theory are only partly corroborated, i.e. algal biomass actually increases with increasing productivity, although the grazer community is released from predation. In more species-rich and complex systems, e.g temperate lakes with three functionally important links, such adaptations are likely to be even more important, and, consequently, the observable effects of trophic interactions from top predators on lower trophic levels even more obscured.     

OPINION Manipulating lake community structure: where do we go from here?

SUMMARY. 1 More than 10 years experience with whole lake pelagic manipulation has suggested some general trends applicable to all freshwater pelagic communities and some specific trends related to lake depth.
2 Among the general trends is the observation that the trophic cascade is strongly damped. This means that changes in phytoplankton biomass can be assured only when the fish community is strongly manipulated.
3 Among the depth related trends is the observation that in shallow lakes, changes in fish community structure are more likely to have cascading impacts on phytoplankton than are changes in deep lakes.
4 In shallow lakes, fish removal frequently results in decreased turbidity which is associated with the development of dense macrophyte populations and significant reductions of algal standing stocks. The mechanisms involve: increased grazing by zooplankton, the removal of fish induced bioturbation and nutrient recycling, and direct and indirect macrophyte effects (shading, zooplankton refuges and competition for nutrients).
5 In shallow lakes, where planktivore biomass can be regulated and macrophyte development is acceptable, fish biomanipulalions are likely to result in reduced algal populations and improved water quality.
6 In deep lakes, where macrophytes are not as important, long-term effects of fish manipulations are strongly dependent upon the probability of non-grazable algal bloom development. This is determined by many factors (chemical, physical and grazer related) which modify the impact that grazers have on phytoplankton biomass.
7 In deep lakes, successful fish biomanipulations may only be effective when chemical and physical factors are altered to produce algal species compositions that permit strong top-down control of prey by predators.     

The effect of large macroinvertebrate herbivores on sessile epibenthos in a mountain stream

Studies of benthic invertebrates in lakes and streams suggest thatlarge-bodied herbivores are more efficient grazers than smallerones. In order to assess the effect of larger herbivores on smallergrazing invertebrates, the presence of dominant grazer taxa wasmanipulated in streamside troughs in a first order temperaterainforest stream in British Columbia. The presence of mayflies(Ameletus sp.) and tailed frog tadpoles (Ascaphustruei) reduced both algal biomass and the abundance of herbivorouschironomids (Orthocladiinae) on ceramic tiles. This confirms thatlarge mobile grazers in streams have a negative effect on smallersessile invertebrate grazers either through resource competition ordirect consumption (predation).     

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.     

High nutrient availability leads to weaker top‐down control of stream periphyton: Compensatory feeding in Ancylus fluviatilis

相似文献   

Ecosystem consequences of diversity depend on food chain length in estuarine vegetation

Biodiversity and food chain length each can strongly influence ecosystem functioning, yet their interactions rarely have been tested. We manipulated grazer diversity in seagrass mesocosms with and without a generalist predator and monitored community development. Changing food chain length altered biodiversity effects: higher grazer diversity enhanced secondary production, epiphyte grazing, and seagrass biomass only with predators present. Conversely, changing diversity altered top‐down control: predator impacts on grazer and seagrass biomass were weaker in mixed‐grazer assemblages. These interactions resulted in part from among‐species trade‐offs between predation resistance and competitive ability. Despite weak impact on grazer abundance at high diversity, predators nevertheless enhanced algal biomass through a behaviourally mediated trophic cascade. Moreover, predators influenced every measured variable except total plant biomass, suggesting that the latter is an insensitive metric of ecosystem functioning. Thus, biodiversity and trophic structure interactively influence ecosystem functioning, and neither factor's impact is predictable in isolation.     

Feedbacks of consumer nutrient recycling on producer biomass and stoichiometry: separating direct and indirect effects

Herbivores can have both direct (consumptive) and indirect (nutrient‐mediated) effects on primary producer biomass and nutrient stoichiometry. Ecological stoichiometry theory predicts that herbivores of contrasting body stoichiometry will differentially remineralize nutrients, resulting in feedbacks on producer stoichiometry. We experimentally separated direct and indirect effects of aquatic vertebrate grazers on periphyton by manipulating grazer abundance and identity in mesocosms, and using grazer exclusion cages to expose periphyton to recycled nutrients in the absence of direct grazing. In experiment 1, we used a catfish with high body phosphorus (low body N:P), Ancistrus triradiatus, to assess consumptive versus nutrient‐mediated effects of grazer density on periphyton. In experiment 2, we compared the nutrient‐mediated effects of grazing by Ancistrus triradiatus and Rana palmipes, a tadpole with low body phosphorus and high body N:P. In experiment 1, we found that increasing catfish density led to lower biomass and particulate nutrients in periphyton through direct consumptive effects, but that nutrient‐mediated indirect effects enhanced periphyton biomass when grazers were experimentally separated from direct contact with periphyton. As predicted by stoichiometry theory, nutrient recycling by this P‐rich grazer tended to increase algal C:P and N:P (although effects were not statistically significant), while their consumptive effects reduced algal C:P and N:P. In experiment 2, grazer identity had strong effects on dissolved water nutrient concentrations, N recycling (measured with a ¹⁵N tracer), and periphyton stoichiometry. In accordance with stoichiometry theory, catfish increased N concentrations and recycling rates leading to higher periphyton N:P, while tadpoles had greater effects on P availability leading to lower periphyton N:P. Our experiments elucidate the importance of both the density and identity of grazers in controlling periphyton biomass and stoichiometry through consumptive and nutrient‐mediated effects, and support the power of ecological stoichiometry theory to predict feedbacks on producer stroichiometry arising from consumer stoichiometry through nutrient recycling.     

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.     

Impacts of grazing intensity and increased precipitation on a grasshopper assemblage (Orthoptera: Acrididae) in a meadow steppe

1. Grasshoppers are dominant herbivores in grassland ecosystems, and many studies have examined how grazing by large herbivores and precipitation patterns individually influence the dynamics of grassland grasshopper assemblages, but their combined effects are largely unknown. 2. In this study, grazing intensities (ungrazed, moderate, and heavy) were manipulated and precipitation (ambient and increased amount of rainfall) altered in a field experiment to test the effects of grazing and altered precipitation on a grasshopper community in a meadow steppe in northeastern China. 3. It was found that grasshopper species richness did not change according to different grazing intensities under ambient precipitation, but was significantly higher (by 38.1%) in moderate grazing intensities under increased precipitation. Grasshopper abundance increased considerably with increasing grazing intensities in ambient precipitation treatments; however, grasshopper abundance in heavy grazing intensities was significantly lower (by 32.9%) than in the other two grazing intensities under increased precipitation. Moreover, the responses of grasshopper abundance to grazing under altered precipitation were species‐specific. 4. Grazing effects on grasshopper species diversity were mediated through the species richness and biomass of grasses (food resources), but the effects on grasshopper abundance were mediated through plant height (vegetation structure) under altered precipitation. 5. These results suggest that appropriate grazing by large herbivores would be considered as beneficial management practices for maintaining grasshopper diversity and abundance under conditions of increased precipitation in grassland ecosystems. Additionally, greater attention should be paid to the population dynamics of different grasshopper species to better understand the responses of grasslands to grazing and altered precipitation.     

Body size and the division of niche space: food and predation differentially shape the distribution of Serengeti grazers

1. Theory predicts that small grazers are regulated by the digestive quality of grass, while large grazers extract sufficient nutrients from low-quality forage and are regulated by its abundance instead. In addition, predation potentially affects populations of small grazers more than large grazers, because predators have difficulty capturing and handling large prey. 2. We analyse the spatial distribution of five grazer species of different body size in relation to gradients of food availability and predation risk. Specifically, we investigate how the quality of grass, the abundance of grass biomass and the associated risks of predation affect the habitat use of small, intermediate and large savanna grazers at a landscape level. 3. Resource selection functions of five mammalian grazer species surveyed over a 21-year period in Serengeti are calculated using logistic regressions. Variables included in the analyses are grass nitrogen, rainfall, topographic wetness index, woody cover, drainage lines, landscape curvature, water and human habitation. Structural equation modelling (SEM) is used to aggregate predictor variables into 'composites' representing food quality, food abundance and predation risk. Subsequently, SEM is used to investigate species' habitat use, defined as their recurrence in 5 × 5 km cells across repeated censuses. 4. The distribution of small grazers is constrained by predation and food quality, whereas the distribution of large grazers is relatively unconstrained. The distribution of the largest grazer (African buffalo) is primarily associated with forage abundance but not predation risk, while the distributions of the smallest grazers (Thomson's gazelle and Grant's gazelle) are associated with high grass quality and negatively with the risk of predation. The distributions of intermediate sized grazers (Coke's hartebeest and topi) suggest they optimize access to grass biomass of sufficient quality in relatively predator-safe areas. 5. The results illustrate how top-down (vegetation-mediated predation risk) and bottom-up factors (biomass and nutrient content of vegetation) predictably contribute to the division of niche space for herbivores that vary in body size. Furthermore, diverse grazing assemblages are composed of herbivores of many body sizes (rather than similar body sizes), because these herbivores best exploit the resources of different habitat types.     

A model approach to planktonic stoichiometry and consumer-resource stability

1. The model explores stoichiometric feedback effects between an alga and a grazer ( Daphnia ) in a simplified chemostat-type system in stagnant conditions or with fixed dilution rate.
2. When running the model with fixed stoichiometry and P-sufficient food, the grazer with highest requirements for phosphorus (P) will exert the most efficient control of algal biomass owing to more P being allocated to zooplankton biomass and less P recycled.
3. When including potential P-limitation of the grazer, the grazer with high P requirements (high P : C ratio) will be the least efficient grazer in a system with fluctuating and temporarily low P : C ratio in algae ( Q _a).
4. Qualitatively deficient food will yield decreased growth efficiency in zooplankton. As Q _a decreases, the grazer isocline for zero net growth is shifted upwards, and the required algal biomass for positive growth increases. There may then be a critical level of Q _a below which the grazer with high P : C suffers negative population growth regardless of algal biomass. In cases with low minimum Q _a and a P-demanding grazer, this may cause the system to enter an irreversible stage with high biomass of P-deficient phytoplankton which do not support zooplankton growth.
5. Cumulative primary production for scenarios with continuous P input is, in general, higher the more Q _a is allowed to drop below saturation values, and highest when this is combined with a grazer with a high P : C ratio. The lower growth rate of P-deficient phytoplankton was compensated for by reduced success of the P-limited grazer, yielding low grazing pressure and resulting in larger phytoplankton biomass.     

Calanoid copepod grazing on phytoplankton: seasonal experiments on natural communities

Calanoid copepods are major components of most lacustrine ecosystems and their grazing activities may influence both phytoplankton biomass and species composition. To assess this we conducted four seasonal, in situ, grazing experiments in eutrophic Lake Rotomanuka, New Zealand. Ambient concentrations of late stage copepodites and adults of calanoid copepods (predominantly Calamoecia lucasi, but with small numbers of Boeckella delicata) were allowed to feed for nine days on natural phytoplankton assemblages suspended in the lake within 1160 litre polyethylene enclosures. The copepods reduced the total phytoplankton biomass of the dominant species in all experiments but were most effective in summer (the time of highest grazer biomass) followed by spring and autumn. In response to grazing pressure the density of individual algal species showed either no change or a decline. There were no taxa which increased in density in the presence of the copepods. The calanoid copepods suppressed the smallest phytoplankton species (especially those with GALD (Greatest Axial Linear Dimension) < µm) and there appeared to be no selection of algae on the basis of biovolume. Algal taxa which showed strong declines in abundance in the presence of the copepods include Cyclotella stelligera, Coelastrum spp., Trachelomonas spp., Cryptomonas spp., and Mallomonas akrokomos. Calanoid copepods are considered important grazers of phytoplankton biomass in this lake. The study supports the view that high phytoplankton:zooplankton biomass ratios and large average algal sizes characteristic of New Zealand lake plankton may, at least partly, be caused by year round grazing pressure on small algae shifting the competitive balance in favour of larger algal species.     

Relative strengths of benthic algal nutrient and grazer limitation along a lake productivity gradient

The relative effects of nutrients and herbivores on primary producers are rarely compared across ecosystems that vary in potential primary productivity. Furthermore, proposed mechanisms to explain such patterns remain understudied. Here, I examine the strength of nutrient and grazer (herbivore) limitation (i.e., the extent to which producers’ growth is limited by insufficient nutrient supply or herbivory) of benthic algae across 13 southwest Michigan lakes that vary widely in productivity (i.e., resource supply). I compare the observed patterns of algal limitation and species composition to those predicted by two simple models: one that includes multiple species and species’ traits (the food-web model) and one that includes no variation in species or traits (the food-chain model). Species in the food-web model are assumed to display a tradeoff between resource competitive ability and resistance to herbivory (the “keystone predator” tradeoff). Among these lakes, benthic algal nutrient limitation was positive and declined significantly along a lake N:P gradient. In contrast, grazer limitation was negative and was not significantly related to any of the measured lake productivity variables. Negative grazer limitation indicated that the removal of grazers caused unexpected declines in algal biomass, which were potentially due to indirect, positive effects (e.g., nutrient recycling) of grazers. Nutrient limitation was significantly stronger than grazer limitation across lakes, which was more consistent with the food-web versus food-chain model. Changes in algal composition were also broadly consistent with predictions of the food-web model in that vulnerable, superior nutrient competitors dominated in low productivity lakes and more grazer-resistant species were observed in high productivity lakes. In general, these results point to the importance of examining limiting factors across systems and the consideration of key species’ traits when predicting and interpreting patterns.     

POSITIVE FEEDBACK AND THE DEVELOPMENT AND PERSISTENCE OF ECOSYSTEM DISRUPTIVE ALGAL BLOOMS1

Harmful algal blooms (HABs) have occurred with increasing frequency in recent years with eutrophication and other anthropogenic alterations of coastal ecosystems. Many of these blooms severely alter or degrade ecosystem function, and are referred to here as ecosystem disruptive algal blooms (EDABs). These blooms are often caused by toxic or unpalatable species that decrease grazing rates by planktonic and benthic herbivores, and thereby disrupt the transfer of nutrients and energy to higher trophic levels, and decrease nutrient recycling. Many factors, such as nutrient availability and herbivore grazing have been proposed to separately influence EDAB dynamics, but interactions among these factors have rarely been considered. Here we discuss positive feedback interactions among nutrient availability, herbivore grazing, and nutrient regeneration, which have the potential to substantially influence the dynamics of EDAB events. The positive feedbacks result from a reduction of grazing rates on EDAB species caused by toxicity or unpalatability of these algae, which promotes the proliferation of the EDAB species. The decreased rates also lower grazer‐mediated recycling of nutrients and thereby decrease nutrient availability. Since many EDAB species are well‐adapted to nutrient‐stressed environments and many exhibit increased toxin production and toxicity under nutrient limitation, positive feedbacks are established which can greatly increase the rate of bloom development and the adverse effects on the ecosystem. An understanding of how these feedbacks interact with other regulating factors, such as benthic/pelagic nutrient coupling, physical forcing, and life cycles of EDAB species provides a substantial future challenge.     

The zooplankton-phytoplankton interface in lakes of contrasting trophic status: an experimental comparison

We report here the results of an experimental study designed to compare algal responses to short-term manipulations of zooplankton in three California lakes which encompass a broad range of productivity (ultra-oligotrophic Lake Tahoe, mesotrophic Castle Lake, and strongly eutrophic Clear Lake). To assess the potential strength of grazing in each lake, we evaluated algal responses to a 16-fold range of zooplankton biomass. To better compare algal responses among lakes, we determined algal responses to grazing by a common grazer (Daphnia sp.) over a range ofDaphnia densities from 1 to 16 animals per liter. Effects of both ambient grazers andDaphnia were strong in Castle Lake. However, neither ambient zooplankton norDaphnia had much impact on phytoplankton in Clear Lake. In Lake Tahoe, no grazing impacts could be demonstrated for the ambient zooplankton butDaphnia grazing had dramatic effects. These results indicate weak coupling between phytoplankton and zooplankton in Clear Lake and Lake Tahoe, two lakes which lie near opposite extremes of lake trophic status for most lakes. These observations, along with work reported by other researchers, suggest that linkages between zooplankton and phytoplankton may be weak in lakes with either extremely low or high productivity. Biomanipulation approaches to recover hypereutrophic lakes which aim only to alter zooplankton size structure may be less effective if algal communities are dominated by large, inedible phytoplankton taxa.     

Modelling ungulate dependence on higher quality forage under large trees in African savannahs

In African savannahs, large trees improve grass quality, particularly in dry and nutrient poor areas. Enhanced below-canopy grass nutrients, such as nitrogen and phosphorus contents should therefore attract and benefit grazers. To predict whether ungulates really need these forage quality islands we focused on four grazer species, i.e., zebra, buffalo, wildebeest, and warthog, differing in body size and digestive system. We confronted literature estimations of their feeding requirements with forage availability and quality, observed in three South African savannah systems, through linear modelling. The model predicted the proportion of below-canopy grass that grazers should include in their diet to meet their nutritional requirements.During the wet season, the model predicted that all animals could satisfy their daily nutrient requirements when feeding on a combination of below- and outside-canopy grasses. However, wildebeest, having relatively high nutrient demands, could meet their nutrient requirements only by feeding almost exclusively below canopies.During the dry season, all animals could gain almost twice as much digestible protein when feeding on below – compared to outside-canopy forage. Nonetheless, only warthogs could satisfy their nutrient requirements – when feeding almost exclusively on below-canopy grasses. The other ungulate species could not meet their phosphorus demands by feeding at either site without exceeding their maximum fibre intake, indicating the unfavourable conditions during the dry season.We conclude that grazing ungulates, particularly warthog, zebra, and buffalo, actually depend on the available below-canopy grass resources. Our model therefore helps to quantify the importance of higher quality forage patches beneath savannah trees. The composition of grazer communities depending on below-canopy grasses can be anticipated if grazer food requirements and the abundance of large trees in savannahs are known. The model suggests that the conservation of large single-standing trees in savannahs is crucial for maintenance of locally grazing herbivores.     

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.     

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.     

Effects of avian grazing on the algal community and small invertebrates in the rocky intertidal zone

In this study of a rocky intertidal habitat in northern Japan, feeding by avian consumers had significant effects on algal assemblages and small herbivorous invertebrates. The effects of the birds on algae were different from those of invertebrate grazers such as urchins and gastropods. The abundance of the dominant algal species decreased during the grazing period, increased again after the grazing period, and indirectly affected algal species richness and evenness. Avian grazing also decreased the density of tube-dwelling amphipods on the dominant alga, but did not change the density of mobile and free-living isopods. These results suggest that avian grazers may act as habitat modifiers rather than exploitative competitors for the small herbivorous crustaceans. Avian herbivores consumed only the upper parts of large algal fronds, apparently reducing the amount of suitable microhabitat for the small herbivorous crustaceans, which are subject to a variety of physical or biological stress. Thus, avian herbivores function as ecosystem engineers, regulating community structure in a manner different to invertebrate herbivores in rocky intertidal habitats.