Experimental Removal and Recovery of Subtidal Grazers Highlights the Importance of Functional Redundancy and Temporal Context

The extent to which different grazers are functionally redundant has strong implications for the maintenance of community structure and function. Grazing by red urchins (Strongylocentrotus franciscanus) on temperate rocky reefs can initiate a switch from invertebrate or macroalgal dominance to an algal crust state, but can also cause increases in the density of molluscan mesograzers. In this study, we tested the hypothesis that red urchins and lined chitons (Tonicella spp.) are redundant in the maintenance of available space, defined as encrusting algae and bare rock. In a factorial field experiment replicated at three sites, we reduced the densities of urchins and chitons on subtidal rock walls for nine months. The effects of grazers were interpreted in the context of natural temporal variation by monitoring the benthic community one year before, during, and after grazer removal. The removal of each grazer in isolation had no effect on the epilithic community, but the removal of both grazers caused an increase in sessile invertebrates. The increase was due primarily to clonal ascidians, which displayed a large (∼75%) relative increase in response to the removal of both grazers. However, the observed non-additive responses to grazer removal were temporary and smaller than seasonal fluctuations. Our data demonstrate that urchins and chitons can be redundant in the maintenance of available space, and highlight the value of drawing conclusions from experimental manipulations within an extended temporal context.     

Consistency in a marine algal‐grazer interaction over multiple scales

Coralline algae are conspicuous members of many marine assemblages, especially those characterized by intense grazing pressure. We explored whether articulated species, especially Corallina vancouveriensis, depend on grazing invertebrates to both establish and flourish in an exposed rocky intertidal setting, and whether this plant–grazer relationship varied over more than three orders of magnitude (≈100–>300,000 μm). Three experimental manipulations, supplemented by observations on recruitment, demonstrated that (i) C. vancouveriensis failed to recover rapidly from disturbed areas when grazers were experimentally excluded; (ii) recruitment occurred in the presence of grazers; (iii) increasing surface texture of molded surfaces enhanced coralline recruitment more when grazers were present; and (iv) settlement occurred predominately in microtopographical low areas of a molded surface, whereas a competitively superior fleshy red alga tended to recruit to high areas. These results confirm that coralline algal establishment and persistence are enhanced by grazers and reveal that this relationship is consistent over a range of biologically relevant scales.     

The effect of grazer size manipulation on periphyton communities

Summary We examined the effect of grazer size on periphyton biomass, size structure, and species compostion by removing the largest invertebrate grazers on artificial macrophytes planted in the littoral of Lake Memphremagog (Que-Vt). A series of exclosures with increasingly fine mesh prevented colonization by large invertebrates but allowed in smaller grazers. Oligochaetes, chironomids, and cladocerans effectively replaced snails so that total grazer biomass in the various treatments was not significantly different from the controls. With one exception, algal biomass, measured as chlorophyll a, did not differ significantly among the various treatments. However algal size and taxonomy were affected because the dominance of large blue-green colonies was apperantly related to the presence of large grazers. The results of the size manipulations were qualitatively similar to those induced in phytoplankton communities by size selective zooplankton grazing and are consistent with models based on general allometric equations.Contribution 181 of the Lake Memphremagog Project, Limnology Research Centre     

Effects of Chiton granosus (Frembly, 1827) and other molluscan grazers on algal succession in wave exposed mid-intertidal rocky shores of central Chile

Molluscan grazers can have important effects on the abundance, colonization rates, and successional pathways of algal assemblages and the entire intertidal community. In general, early successional algae are more readily consumed than corticated algae and kelps, which usually get established later in the community succession. To generalize, however, the effect of different grazers on algal assemblages must be examined on different coasts and under different scenarios. This information could help us understand the mechanisms of ecosystem processes and situations in which general models do not apply. Along the coast of Chile, humans harvest large keyhole limpets, which seem to be the only invertebrate grazers capable of controlling the dominant corticated alga Mazzaella laminarioides, a canopy-forming species that can cover extensive areas of the mid intertidal zone. In this scenario, where large limpets are harvested, the overall effects of the diverse molluscan assemblage of limpets, chitons and snails on algal succession and on corticated algae in particular are not clear. We conducted a 26-month-long experiment to evaluate the effects of molluscan grazers on mid-intertidal algal succession and to isolate the effects of Chiton granosus, the most conspicuous member of the assemblage at these tidal elevations. At sites heavily impacted by humans the molluscan grazer assemblage had strong negative effects on colonization and abundance of green algae such as ulvoids and Blidingia minima. In doing so, the grazer assemblage had a strong negative indirect effect on the establishments of chironomid fly larvae, which were only observed on green algal mats and rarely on bare rock. No significant effects were detected on epilithic microalgae, and effects on sessile invertebrates were highly variable over space and time. C. granosus also had significant negative effects on green algae but did not account for the total grazing pressure exerted by the guild. Limited foraging excursions (ca. 35 cm) from refuges and moderate site (crevice) fidelity in this species may contribute to the patchiness in green algal distribution observed in the field. Nearly 13 months after rock surface were experimentally cleared, M. laminarioides appeared in all experimental plots, but increased over three times faster in enclosures containing C. granosus than in exclosures plots or controls, suggesting that moderate levels of herbivory could actually facilitate the establishment of this alga in the succession and that the green algal cover found in the absence of grazers may delay its establishment.     

Algal propagule banks modify competition, consumer and resource control on Baltic rocky shores

We hypothesized that supply from macroalgal propagule banks may influence the relative abundance of annual and perennial algae and that this may alter the effects of grazers and nutrients on species composition. In a factorial field experiment in the Baltic Sea littoral system we tested the effects of manipulating propagule banks, the abundance of crustacean and gastropod grazers, and nutrient supply on recruitment and growth of macroalgae over a year. Moreover, we determined seasonal patterns of macroalgal propagule dispersal at the experimental site and quantified algal abundance and recruitment at 25 locations throughout the Baltic Sea. Experimental manipulations had minor effects on adults of the dominating perennial alga, Fucus vesiculosus. Instead, we found that species composition was determined by processes operating at early life stages. Propagule supply from a propagule bank strongly favored the fast-growing annual alga Enteromorpha spp. which then blocked settlement and recruitment of Fucus. Grazers reduced the abundance of annual algae and indirectly favored Fucus recruitment. There was an apparent trade-off between gains from the propagule bank and losses to herbivory in five of seven colonizing species. Nutrient enrichment overrode grazer control of annual algae and accelerated the decline of Fucus only when annual algae had already achieved high densities through the propagule bank. Corroborating the experimental findings, field surveys across the Baltic showed that Fucus recruit densities can be predicted from the cover of annual algae during the period of Fucus reproduction and settlement. Recruitment inhibition by annual algae, which is driven by the abundance of annuals in the propagule bank, increasing nutrient levels, and declining consumer control, is suggested as a mechanistic explanation of the current decline of perennial algae in the Baltic Sea.     

Loss processes influencing growth of planktonic bacterial populations in Lake Constance

The fate of bacterial production in a deep mesotrophic lakehas been investigated on the basis of a multiple approach combiningfield and experimental studies. Bacterial production gains asestimated by measurements of methyl-[³H]thymidine incorporationinto macromolecules appeared to be more or less balanced bycorresponding grazing losses. This was suggested by consideringabundance and grazing potential of presumed grazers as wellas by following in time the binding of labelled bacteria bylarger sized organisms. Moreover, strong increases in bacterialnet growth were always observed after removal of grazers fromthe water samples. As was indicated by independent approaches,phagotrophic microflagellates were mainly responsible for theobserved grazing losses. Support for the importance of alternativeloss processes such as sinking or autolysis was given neitherby field observations nor by long-term expenments with exclusionof grazers. It is concluded that phagotrophic microflagellatesmay be regarded as important functional components of planktoniccommunities in freshwater lakes.     

Forage patch use by grazing herbivores in a South African grazing ecosystem

Understanding how different herbivores make forage patch use choices explains how they maintain an adequate nutritional status, which is important for effective conservation management of grazing ecosystems. Using telemetry data, we investigated nonruminant zebra (Equus burchelli) and ruminant red hartebeest (Alcelaphus buselaphus subspecies camaa), use of burnt patches in a landscape mosaic of nutrient-poor, old grassland interspersed with young, recently burnt, nutrient-rich grass patches. The Mkambati Nature Reserve landscape on the east coast of South Africa provided large grazers with a challenge in finding and using appropriate patches in which to forage to meet their nutritional requirements. In Mkambati, grassland fires, mostly ignited by poachers, induce regrowth of young nutrient-rich grass, which subsequently attract grazers. We tested if the study animals foraged more in burnt patches than in the unburned grassland and whether burnt patch use was related to the distance to the previously visited burnt patch, burnt patch size, burnt patch age, and distance to areas with high poaching risk using MANOVA. In general, zebra moved faster than red hartebeest, and both species moved faster in unburnt grassland than in burnt patches. Red hartebeest and zebra patch selection were influenced by interpatch distance, patch age, patch size, and poaching risk. A limited set of intrinsic traits, i.e., body mass, digestion strategy, and muzzle width, yielded different patch use rules for the two species. Large ungulates patch use behaviour varied among species and across conditions and was influenced by anthropogenic impacts such as poaching and changed fire regimes. This could potentially affect biodiversity negatively and needs to be factored into management of conservation areas.     

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.     

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.     

Grazing on toxic Alexandrium fundyense resting cysts and vegetative cells by the eastern oyster (Crassostrea virginica)

In laboratory experiments, oysters (Crassostrea virginica) were fed Alexandrium fundyense (strain CB501) vegetative cells or resting cysts (from strains CB501 and GMT25) produced from laboratory cultures. The toxicity per cyst was 1.7 pg STXequiv/cyst and for vegetative cells 3.9 pg STXequiv/cell. The toxic, resting cysts and vegetative cells were removed from suspension in the experimental containers within about 4 h. Oysters fed toxic vegetative cells digested 72% of cells ingested, and 28% survived gut passage by forming temporary cysts. Toxin levels of oysters fed vegetative cells averaged 27 μg STXequiv/100 g meat. Resting cysts added to the experimental containers adhered to the walls so that only 40% of the cysts added were available to the oysters during the experiment. Of the cysts that were ingested, approximately 59% were digested, and oysters accumulated toxins (an average of 1.2 μg STXequiv/100 g meat), showing that consumption of resting cysts can cause toxicity in oysters. Direct consumption of resting cysts, thus, may explain shellfish toxicity in areas without known blooms, but with toxic resting cysts in the sediment. These results suggest a possible role of toxic cysts in mediating time-lags between surface blooms and appearance of toxicity in benthic grazers, and the possible role of benthic grazers in controlling seed populations, except in anoxic areas, which can serve as cyst “refuges” from grazing mortality.     

Increasing fish taxonomic and functional richness affects ecosystem properties of small headwater prairie streams

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The effects of grazing by gastropods and physical factors on the upper limits of distribution of intertidal macroalgae

Summary The cover of foliose algae is sparse to non-existent above a low-level algal zone on many shores in N.S.W., except in rock-pools. Above this algal zone, encrusting algae, mostly Hildenbrandia prototypus, occupy most of the primary substratum on sheltered shores. Experimental manipulations at midtidal levels were used to test hypotheses about the effects of grazing by molluses and of physical factors during low tide on this pattern of algal community structure.Fences and cages were used to exclude grazers: molluscs grazed under roofs and in open areas. Cages and roofs provided shade, and decreased the harshness of the environment during low tide: fences and open areas had the normal environmental regime.In the absence of grazers, rapid colonization of Ulva and slower colonization by other foliose algae occurred in all experimental areas. The rate of colonization by Ulva sporelings was initially retarded on existing encrusting algae, but after a few months, cover of Ulva equalled that on cleared rock.Most species of algae only grew to maturity inside cages, and remained as a turf of sporelings inside fences. No foliose algae grew to a visible size in open, grazed areas. Grazing thus prevents the establishment of foliose algae above their normal upper limit on the shore, but the effects of physical factors during low tide prevent the growth of algae which become established when grazers are removed. Physical factors thus limit the abundance of foliose algae at mid-tidal levels.The recolonization of cleared areas by Hildenbrandia was not affected by the presence of a turf of sporelings, nor by the shade cast by roofs, but was retarded in cages where mature algae formed a canopy. Even under such a canopy, Hildenbrandia eventually covered as much primary substratum as in open, grazed areas. This encrusting alga is able to escape from the effects of grazing by having a tough thallus, and by its vegetative growth which allows individual plants to cover a lot of substratum, and by the tendency for new individuals to start growing from small cracks and pits in the rock, which are apparently inaccessible to the grazers.Mature foliose algae are removed from the substratum by waves, and many individual plants died during periods of hot weather. Sporelings in a turf were eliminated, after experimental fences were removed, by the combined effects of macroalgal grazers, which invaded the areas, and microalgal grarers which ate the turt from the edges inwards.The results obtained here are discussed with respect to other studies on limits to distribution of intertidal macroalgae, and the role of grazing in the diversity and structure of intertidal algal communities. Some problems of these experimental treatments are also discussed.     

Interactions between planktonic microalgae and protozoan grazers

For an algal bloom to develop, the growth rate of the bloom-forming species must exceed the sum of all loss processes. Among these loss processes, grazing is generally believed to be one of the more important factors. Based on numerous field studies, it is now recognized that microzooplankton are dominant consumers of phytoplankton in both open ocean and coastal waters. Heterotrophic protists, a major component of microzooplankton communities, constitute a vast complex of diverse feeding strategies and behavior which allow them access to even the larger phytoplankton species. A number of laboratory studies have shown the capability of different protistan species to feed and grow on bloom-forming algal species. Because of short generation times, their ability for fast reaction to short-term variation in food conditions enables phagotrophic protists to fulfill the function of a heterotrophic buffer, which might balance the flow of matter in case of phytoplankton blooms. The importance of grazing as a control of microalgae becomes most apparent by its failure; if community grazing controls initial stages of bloom development, there simply is no bloom. However, if a certain algal species is difficult to graze, e.g. due to specific defense mechanisms, reduced grazing pressure will certainly favor bloom development. The present contribution will provide a general overview on the interactions between planktonic microalgae and protozoan grazers with special emphasis on species-specific interactions and algal defense strategies against protozoan grazers.     

Do small-scale exclosure/enclosure experiments predict the effects of large-scale extirpation of freshwater migratory fauna?

A variety of theoretical and empirical studies indicate that the abilities of small-scale experiments to predict responses to large-scale perturbations vary. Small-scale experiments often do not predict the directions of large-scale responses, and relatively few empirical studies have examined whether small-scale experiments predict the magnitudes of large-scale responses. Here we present an empirical example of small-scale manipulations predicting not only the directions but also the magnitudes of the effects of whole-catchment, decades-long decimation of migratory freshwater shrimp populations. In streams of Puerto Rico (USA), we used arena sizes of < 2 m² in 1- to 4-week exclosure/enclosure experiments. Effects of small-scale experiments largely matched those of large-scale shrimp loss above dams for a variety of response variables (abiotic and biotic factors including epilithic fine sediments, algae and organic matter, and invertebrate grazers, detritivores, and predators). The results of our extrapolation contrast with studies of small- versus large-scale perturbations in the temperate zone. Our findings are likely explained by: a set of response variables that are more dominated by within-patch processes than exchange processes, an experimental manipulation that encompassed the characteristic scales of response variables, our use of open arenas lacking cage artifacts, and/or our combination of two distinct experimental approaches (exclosures and enclosures). Based on our study design, we suggest that extrapolation across experimental scales can be greatly enhanced by embedding open arenas within large-scale conditions that represent all treatment levels. Electronic Supplementary Material Supplementary material is available to authorised users in the online version of this article at .     

Iron-Alum Aceto-Carmine Staining for Chromosomes and Other Anatomical Features of Rhodophyceae

The chromosomes, certain intracellular structures and gross anatomical details of many red algae, which, as a class, have proved technically difficult material, can be demonstrated by staining with aceto-carmine after a mordant bath of iron alum. Acetic-alcohol mixtures are used as nuclear fixatives and formalin-acetic-alcohol and other similar fluids for preservation of anatomical features. The tougher more cartilaginous thalli of some species can be softened, if squashes are desired, by prolonging fixation (24-48 hr.) in acetic alcohol and subsequent washing. The fixatives are washed out of the material before the latter is transferred to 0.5-5.0% ferric ammonium sulphate, the concentration of which may be altered according to the material. Excess mordant is removed by washing and the material stained in Belling's aceto-carmine containing a trace of ferric acetate as a “ripener”. The degree of heating before covering is critical as it controls the quality of the staining. Squashing must be very thorough to spread the chromosomes which are usually very small but only slight controlled pressure is necessary when diffuse structures such as carposporophytes, nemathecia or medullary filaments are being demonstrated. Paraffin sections mounted on slides can also be stained by this method.     

Interspecific Competition for Shelters in Territorial and Gregarious Intertidal Grazers: Consequences for Individual Behaviour

Experiments have shown that interspecific interactions within consumer guilds can alter patterns of distribution, abundance and size of species. Plastic behavioural responses can be modulated by agonistic interactions. In many cases, consumers compete for space and shelters, and these interactions change the manner in which they exploit food. This study investigates the consequences of competition in the spatial and temporal organization of behaviour of intertidal grazers, which share algal resources and the use of rock crevices while resting, but exhibit different body sizes, spatial behaviour and foraging modes. We evaluate interaction strength between small gregarious Siphonaria lessoni and the larger territorial keyhole limpet Fissurella crassa and between S. lessoni and the medium-size gregarious chiton Chiton granosus. Using field manipulations and artificial arenas in the laboratory, we tested whether the use of crevices, micro-spatial distribution and activity are modified by the density of conspecifics and the presence of heterospecifics. Our results show that small-scale spatial segregation observed in the field between S. lessoni and C. granosus result from species-specific differences in habitat use. In turn, we found evidence that spatial segregation between F. crassa and S. lessoni results from highly asymmetric interference competition in the use of shelters. The presence of F. crassa reduced the use of crevices and growth rates of S. lessoni. Effects on growth rates are assumed to result from exposure to harsh environmental conditions rather than food limitation. Thus, neither gregarious behaviour nor differences in activity were sufficient to prevent competition with the larger grazer. Our study illustrates the importance of competition for shelters, which results in behavioural changes of the smaller-sized species, and how these plastic responses can translate into differences in growth rates. Use of shelters can thus be modulated by environmental conditions in a species-specific as well as an interactive manner within consumers’ guilds.     

Genetic engineering, welfare, and accountability

Comments on the implications of genetic engineering for animal welfare. Welfare problems associated with techniques used to achieve genetic changes; Detrimental effects of genetic modifications to welfare; Modification of farm animals for biomedical products. Implications of genetic engineering for animal welfare are changing rapidly and need to be reviewed regularly. They include the welfare problems associated with techniques used to achieve genetic changes, which are similar to problems of other experimental approaches; these should be considered carefully, especially where techniques are used on a routine basis. When it comes to the genetic modifications themselves, some are detrimental to welfare, some are neutral, and some are beneficial; these results include direct effects of the intended change, side effects, and indirect effects. Currently, the two main applications are modification of farm animals for biomedical products--which appears to be largely neutral for welfare--and modification of mice as models for human disease, which results in suffering, often severe suffering. Beneficial applications are rare and still experimental or theoretical. The situation is similar with regard to the use of recombinant hormones and viruses; use of recombinant vaccines has potential for improving welfare, but may raise other ethical problems. Although few, if any, of these concerns are specific to genetic engineering, various factors combine to suggest that particular safeguards are needed in this field. These include the facts that changes can be produced rapidly and repeatedly, and that one of the driving forces behind genetic engineering is commercial exploitation of technology. In general, ethical evaluation still is done on a case-by-case basis, using the limited criteria seen as directly relevant to each case, rather than on a broader framework. There also is little public accountability, whereby the public can have confidence that such evaluation is being carried out properly. Calls for advisory “watchdog ”committees to consider ethical questions on the use of animals are endorsed by this article. Furthermore, it is essential for public confidence in the safeguarding of animal welfare that the procedures of such committees should be well-publicized.     

Taxonomic differences shape the responses of freshwater aerobic anoxygenic phototrophic bacterial communities to light and predation

Aerobic anoxygenic phototrophic (AAP) bacteria are a phylogenetically diverse and ubiquitous group of prokaryotes that use organic matter but can harvest light using bacteriochlorophyll a. Although the factors regulating AAP ecology have long been investigated through field surveys, the few available experimental studies have considered AAPs as a group, thus disregarding the potential differential responses between taxonomically distinct AAP assemblages. Here, we used sequencing of the pufM gene to describe the diversity of AAPs in 10 environmentally distinct temperate lakes, and to investigate the taxonomic responses of AAP communities in these lakes when subjected to similar experimental manipulations of light and predator removal. The studied communities were clearly dominated by Limnohabitans AAP but presented a clear taxonomic segregation between lakes presumably driven by local conditions, which was maintained after experimental manipulations. Predation reduction (but not light exposure) caused significant compositional shifts across most assemblages, but the magnitude of these changes could not be clearly related to changes in bulk AAP abundances or taxonomic richness of AAP assemblages during experiments. Only a few operational taxonomic units, which differed taxonomically between lakes, were found to respond positively during experimental treatments. Our results highlight that different freshwater AAP communities respond differently to similar control mechanisms, highlighting that in‐depth knowledge on AAP diversity is essential to understand the ecology and potential role of these photoheterotrophs.     

Flow rate mediates the competitive influence of a grazing caddisfly on mayflies

The Glossosomatid caddisflies, which are stream-insect grazers, have been found to negatively affect the densities of mayfly grazers through intensive periphyton exploitation. The grazing efficiency of a Glossomatid caddisfly, however, is known to decrease with increasing current velocity. A manipulative field experiment was conducted to test the hypotheses that the presence of Glossosoma sp. decreased the densities of three mayfly grazers, Baetis thermicus, Cinygmula sp. and Paraleptophlebia chocorata, and that the decrease was stronger in slower stream-flow conditions. The experiment followed a two-factorial design with flow conditions and Glossosoma densities as the main factors. The periphyton biomass in conditions of reduced Glossosoma density was greater than in natural Glossosoma densities, but was not affected by flow conditions. In partial contradiction of the experimental hypothesis, the experimental reduction in Glossosoma density resulted in a higher density of Baetis, irrespective of flow conditions. The grazing activity of Glossosoma would not be affected by the flow variations that were employed in the present experiment. The reduction in Glossosoma density resulted in higher densities of Cinygmula and Paraleptophlebia only in fast-flow conditions, a result that did not support the experimental hypotheses. The competitive influence of Glossosoma on these mayflies might be altered by flow conditions causing differences in the resource structure.