Periphyton removal related to phosphorus and grazer biomass level

1. The proliferation of nuisance periphyton in enriched streams may be dependent on the biomass of the grazing macroinvertebrates present. In the present study, the effectiveness of grazer size and biomass in controlling periphyton and the extent to which grazing effectiveness was affected by enrichment level were determined. 2. Two sets of experiments with two caddisfly grazers were conducted in laboratory channels during spring and summer 1995 and 1996. The first set tested the combined effect of phosphorus enrichment and grazing, while the second set tested the effect of variable grazer biomass on periphyton biomass. 3. Grazing reduced periphyton biomass in excess of 80%, compared to ungrazed controls. Grazers were equally effective in controlling filamentous green algae, Stigeoclonium, diatoms and small colonial greens. Near complete removal of periphyton biomass by grazing occurred at even at the lowest grazer biomass level (750 mg m^??2, i.e. approximately one-third of natural levels). 4. Grazing controlled periphyton biomass more than did enrichment with soluble reactive phosphorus (SRP). 5. Grazing rates in the phosphorus-grazing interaction experiments averaged about 6 mg chl a g invertebrate^??1 day^??1, which was similar to past work in these channels and elsewhere, while rates were about five-fold higher in the variable grazer biomass experiments. 6. Simulating effects of SRP and grazing with a calibrated model suggests that higher SRP levels would be necessary to exceed a nuisance periphyton biomass level if grazers were present. However, if grazer biomass was more than 1500 mg m^??2, a nuisance level would probably not be exceeded at any SRP.     

Does an increase in irradiance influence periphyton in a heavily-grazed woodland stream?

Summary Irradiance level and grazer density were manipulated in a factorial design to examine the relative effects of biotic and abiotic factors on periphyton biomass, productivity, and taxonomic structure in a heavily grazed, woodland stream. Irradiance levels were increased from 0.26 to 12.42 mol quanta/m²/d by placing metal halide lamps over the stream. The major grazer in this system was the prosobranch snail Elimia clavaeformis. Its densities were reduced from ca. 750 individuals/m² to near zero by raising platforms off the stream bottom. Experimental treatments were maintained for 48 days. Biomass-specific carbon fixation rates increased significantly in response to higher light levels, indicating that periphyton communities were light-limited at this time of year. However, positive effects of irradiance on areal-specific carbon fixation and biomass were detected only when grazer density was reduced. Basal cells of the chlorophyte Stigeoclonium dominated communities exposed either to low light or high grazing pressure. When light was increased and grazer density reduced, large or upright diatoms became more abundant. Results from this study indicated that limitation of periphyton photosynthesis could be mitigated by increasing the levels of an abiotic resource (light) to this system, but that periphyton biomass was controlled by biotic interactions.     

Experimental Analysis of Grazing by the Mayfly Meridialaris chiloeensis on Different Successional Stages of Stream Periphyton

In this study we determined grazing effects of the South Andean endemic mayfly Meridialaris chiloeensis on periphyton at different stages of successional development. Grazing effects were studied through a two‐factor experimental design (colonization stages X grazer density) in a stream‐side channel in spring and winter. Our results showed an absence of proportionality between grazer density and periphyton decline in response to grazers at low and intermediate levels of periphytic biomass; however, when periphyton biomass was high a direct inverse relationship was observed between post‐grazing biomass and grazer density. The relationship between periphytic algae (chlorophyll a concentration) and periphyton (total periphytic ash‐free dry mass) (C/OM index) was used as an estimation of the autotrophic fraction in the total periphyton matrix. Grazing did not alter the C/OM index indicating that both autotrophic and heterotrophic fractions of the periphyton components were reduced in the same proportion. Ordination of samples using the relative abundance of diatom species showed that herbivore effect was less evident at intermediate and late stage of colonization than at early one. These results support the statement that the outcome of the herbivore‐periphyton interaction may depend on the successional stage of the periphyton community. In spring Fragilaria pinnata relative abundance, on the basis of cell counts, was reduced by grazing and Nitzschia palea was enhanced. In the winter experiment, grazing decreased Achnanthes minutissima relative abundance. (© 2004 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

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.     

Periphyton responds differentially to nutrients recycled in dissolved or faecal pellet form by the snail grazer Theodoxus fluviatilis

1. We aimed to separate the effects of grazers on periphyton via grazing from that of nutrient recycling from their faecal pellets. 2. We set up three different experimental treatments (snails/no snails/faecal pellets) and sampled over 16 days. The ‘snail’ treatment contained a low density (snail biomass c. 14 g^?2) of the gastropod grazer Theodoxus fluviatilis and the ‘faecal pellet’ treatment received the same amount of faecal pellets as were produced in the ‘snail’ treatment. Whereas the ‘faecal pellet’ treatment provided extra nutrients to periphyton from the faeces, the ‘snail’ treatment provided nutrients in the form of both faeces and in excreta. There was also direct grazing on periphyton in the ‘snail’ treatment. The ‘no snail’ was not grazed and received no nutrients in faeces or excreta. 3. We measured periphyton C, N and P content, chlorophyll‐a (chl‐a), primary production, bacterial biomass, bacterial production and bacterial respiratory activity. In the water column we measured dissolved inorganic N and soluble reactive P. 4. Snails increased the amount of dissolved inorganic N in the water. On day 16, the periphyton N : P ratio in the ‘faecal pellet’ treatment was lower, and periphyton P content was higher, than in the other two treatments. N : P ratios decreased over time in the ‘faecal pellet’ treatment. Primary and bacterial production were positively correlated in all treatments. 5. Algal chl‐a and primary production of periphyton per unit area and periphyton chl‐a : C ratios increased over the 16 day in the ‘snail’ treatment, and thus excretion of dissolved N by snails had a stronger positive effect on the periphyton community than N and P in faecal pellets. 6. Our data show that excretion and egestion can have different effects on periphyton, probably because of the higher proportion of dissolved N in excreta and the higher proportion of P recycled in faecal pellets. The relative effect of nutrients recycled in egesta or in excretions, probably depends on the form of nutrient limitation of the periphyton. Further, the different components of the periphyton matrix could react differently to the different forms of nutrient recycling. 7. We conclude that direct grazing effects are less important than nutrient effects when nutrients are limiting and grazing pressure is low. Further, the spatial separation of different grazing effects can lead to differences in periphyton production and nutrient stoichiometry. This might be an explanation for the patchiness of periphyton in nature.     

Periphytic food and predatory crayfish: relative roles in determining snail distribution

Summary In the laboratory and field, we examined how periphyton (food of snails) and predatory crayfish influenced snail distribution in Trout Lake, a permanent, northern Wisconsin lake. Laboratory experiments (with no crayfish) tested the importance of periphyton biomass in determining snail preference among rocks, and among rock, sand, and macrophyte substrates. Among rocks with four different amounts of periphyton, periphyton biomass and the number of Lymnaea emarginata, Physa spp., and Amnicola spp. were positively related. A similar, but non-significant, trend occurred for Helisoma anceps. A field experiment at a site in Trout Lake where predation risk was low confirmed the preference by snails for periphyton covered rocks; more snails colonized rocks with periphyton than rocks without. When given a choice of rock, sand, and macrophytes in the laboratory, L. emarginata preferred high periphyton biomass and rock. Laboratory and field results contrasted with the distribution of snails in Trout Lake; no snails occurred in areas with abundant periphyton-covered rocks, but snails were abundant nearby on scattered rocks with little periphyton. However, where snails were absent, crayfish were abundant (14.5 crayfish-trap^–1-day^–1), and where snails were abundant, crayfish were rare (3.2 crayfish-trap^–1-day^–1), suggesting that crayfish predation reduced snails. The hypothesis that the negative association between snail and periphyton biomass resulted from snail grazing was supported by the results of a field snail enclosure-exclosure experiment (1 m² cages; n=3). All experiments and observations therefore suggest that: 1) crayfish predation is more important than a preference for high periphyton biomass in determining snail distribution in Trout Lake; 2) periphyton biomass is negtively related to snail grazing; and 3) crayfish had a positive indirect effect on periphyton by preying on grazing snails.     

Competition and facilitation within and between a snail and a mayfly larva and the effect on the grazing process

We studied the competitive effects within and between two taxonomically distant freshwater herbivores, a snail and a mayfly, common in Swedish lakes, Lymnaea peregra and Cloeon dipterum, respectively, and their effect on grazing in a laboratory experiment. The experimental set-up consisted of 2-l aquaria, each containing a periphyton covered tile. Intra- and interspecific effects were tested by increasing the density of one species at a time in four different treatments, (1) snails (intraspecific treatment), (2) mayflies (intraspecific treatment), (3) mixed-snails (interspecific treatments, snails kept constant) and (4) mixed-mayflies (interspecific treatments, mayflies kept constant). Intraspecific competition affected both snails and mayflies negatively, i.e. increasing mortality with increasing con-specific density. Furthermore, there was a decrease in snail growth with increasing snail density. In the mixed-species treatments both species changed their microhabitat use indicating interspecific competition. Despite this, we also found a positive effect of mayfly density on snail growth, most likely due to indirect commensalism. No density-dependent effect of grazing on periphyton was found, probably due to interference competition between grazers. However, there was a significant difference in periphyton biomass, due to species composition of grazers. Irrespective of their densities, if they co-existed, the two grazer species decreased the periphyton biomass significantly compared with both single-species treatments. We considered this as a joint action of facilitation and interaction. Our results suggest that competition can be an important structuring factor in macroinvertebrate communities and that species composition can be significant for ecosystem processes within lentic environments.     

How do grazers affect periphyton heterogeneity in streams?

The effects of grazing by stream invertebrates on algal biomass and spatial heterogeneity were tested experimentally in flow-through microcosms with natural substrates (rocks). One experiment tested the effects of fixed densities of three species of grazers (the caddisfly Allomyia sp. and two mayflies, Epeorus deceptivus and Baetis bicaudatus) on periphyton. Baetis was tested with and without chemical cues from fish predators, which reduced grazer foraging activity to levels similar to the less mobile mayfly (Epeorus). Mean algal biomass (chlorophyll a; chl a) was reduced in grazer treatments compared to ungrazed controls, but there were no differences among grazer treatments. Algal heterogeneity (Morisita index) increased with grazer mobility, with the highest heterogeneity occurring in the Baetis-no fish treatment (most mobile grazer) and the lowest in the caddisfly treatment (most sedentary grazer). A second experiment used a three factorial design, and tested whether initial resource distribution (homogeneous vs. heterogeneous), Baetis density (high vs. low) and fish odor (present vs. absent) affected grazer impact on algal resources. Abundances of Baetis and chl a on individual rocks were recorded to explore the mechanisms responsible for the observed distributions of algae. Initial resource heterogeneity was maintained despite being subjected to grazing. Mean chl a was highest in controls, as in experiment I, and effects of Baetis on algal biomass increased with grazer density. There were no fish effects on algal biomass and no effects of grazer density or fish on algal heterogeneity. At the scale of individual rocks Baetis was unselective when food was homogeneously distributed, but chose high-food rocks when it was heterogeneously distributed. Results of these mechanistic experiments showed that Baetis can track resources at the scale of single rocks; and at moderate densities mobile grazers could potentially maintain periphyton distributions observed in natural streams.     

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.     

Stream periphyton response to grazing and changes in phosphorus concentration

Grazing by the large caddisfly larva, Dicosmoecus gilvipes (Trichoptera; Limnephilidae), drastically reduced periphyton biomass in laboratory channels at a current velocity of 20 cm s^–1. Reduction in biomass as chl a and AFDW ranged from 88 to 93% and 82 to 85%, respectively. On average, grazing rate increased with in-channel SRP (soluble reactive phosphorus) content from 6 to 10 µg 1^–1. Grazing rates averaged 25.9–29.3 µg chl a m^–2 d^–1 and 10.8–12.2 µg chl a mg^–1 d^–1 based on area and grazer biomass, respectively, with most variability among treatments being due to the grazing effect. Grazing tended to shift the algal community increasingly to filamentous blue-green algae regardless of enrichment. After three weeks, Phormidium comprised over 61% of the community in grazed treatments but only 35% in ungrazed treatments. The stalked diatom Gomphonema comprised only 4% of the grazed community, but 11% in the three ungrazed channels with similar values for Scenedesmus. A model that includes grazing was calibrated to the data and produced a reasonable expectation of periphyton biomass over a range in SRP concentrations. While the model with constant grazer abundance predicts a gradually increasing grazed biomass as SRP increases, grazer production in natural streams may actually increase to accommodate the increased food production.     

Density-dependent influences on feeding and metabolism in a freshwater snail

We used laboratory experiments to assess the degree of, and the underlying mechanism for, density dependence in the grazing rate of the pulmonate gastropod Physella virgata. Both fecal pellet production and uptake and incorporation of ¹⁴C radioisotopes from labeled periphyton were used as indices of grazing rates. Pronounced density-dependent reductions in grazing rate were observed, especially at densities above 4 snails/-25 cm² periphyton grazing area. Radioisotope experiments also indicated that proportions of ingested ¹⁴C periphyton retained in snail tissue and respired as carbon dioxide increased at higher densities, suggesting that both assimilation efficiency and respiratory costs increase at higher densities. Constant replacement of water in aquaria did not remove density-dependent effects on grazing, suggesting that a dissolved metabolite is not responsible. Experiments where tiles were pre-conditioned with snails grazing at several densities actually stimulated grazing in subsequently added snails, suggesting that substrate-borne cues are also not responsible for density-dependent reductions in grazing rate. Behavioral inferference (in the form of shell-shaking after contacts with other snails) did, however, increase at higher densities, and may be partially reponsible for depressed grazing rates.     

Indirect effects of a parasite on a benthic community: an experiment with trematodes, snails and periphyton

1. Few studies have directly addressed the role played by parasites in the structure and function of ecosystems. Parasites influence the behaviour, reproduction and overall fitness of their hosts, but have been usually overlooked in community and ecosystem‐level studies. We investigated the effects of trematode parasites on snail–periphyton interactions. 2. Physa  acuta (Gastropoda: Pulmonata) snails infected with the trematode Posthodiplostomum minimum (often >30% of within‐shell biomass) grazed more rapidly than uninfected snails. Trematode effects on snail grazing indirectly affected the standing stock and community structure of periphyton. Populations of snails with 50% infected individuals reduced algal biomass by 20% more than populations with lesser (10% or 0%) infection rates. 3. The alga Cladophora glomerata dominated periphyton communities grazed by snail populations with 50% infection rates, whereas diatoms and blue–green algal taxa dominated when grazed by snail populations with lower infection rates. 4. Thus, trematodes indirectly affected periphyton communities by altering host snail behaviour, a trait‐mediated indirect effect. These results indicate that trematodes can indirectly influence benthic community structure beyond simple population fitness, with possible related effects on ecosystem function.     

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

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Do submerged aquatic plants influence their periphyton to enhance the growth and reproduction of invertebrate mutualists?

It has been suggested that submerged aquatic plants can influence the nutritional quality of the periphyton which grows on their surfaces, making it more nutritious for grazing invertebrates, particularly snails. In return, these grazers might preferentially feed on the periphyton and clear the plants of a potential competitor, with the plants and grazers both gaining from this mutualistic relationship. A highly replicated experiment was conducted, in which the nature of the plant (isoetid and elodeid types compared with similar-shaped inert substrata), the nutrient loading, and the influence of periphyton grazers (the bladder snail, Physa fontinalis) of similar size and history were controlled. Plant growth and survival significantly increased in the presence of the periphyton grazer. Whilst the presence of the grazers had the largest influence on periphyton abundance, nutrient availability and plant type also had effects. Plant type had little influence on the nutritional quality of the periphyton measured as carbohydrate, protein and C:N. Effects of treatment on snail growth, and the timing and extent of snail reproduction disappeared when they were compared with the quantity of periphyton available. There was no evidence of enhanced grazer success in the presence of the live plants compared with inert substrata. Although submerged plants affect the growth and reproduction of the grazers which feed on their surfaces, through differences in the amount of periphyton which grows there, we found no evidence that they manipulate the periphyton to encourage such grazers. Received: 1 September 1998 / Accepted: 12 May 1999     

Empirical analysis of the removal rate of periphyton by grazers

To establish a general model for the removal rate of periphyton by grazers, we identified 27 publications in which removal rates could be estimated from grazer enclosure or exclosure experiments. When all the measurements obtained under different experimental conditions were extracted, these publications provided 107 data points. Multiple regression of these data showed that periphyton removal rate increased significantly with grazer body mass and food availability, and decreased with grazer crowding. Grazer body mass explained 65% of the variation, while crowding and food availability explained 7 and 6% respectively. Except for the significantly lower removal rate of amphibians, neither taxon of the grazer nor algal composition significantly affected removal rate. Experiments in the laboratory and in outdoor channels tended to give higher removal rates than experiments performed in streams or lakes. A comparison with previous allometric equations predicting the ingestion rate of other invertebrate guilds, and with experiments in which periphyton ingestion rate was measured as incorporation of labelled food, indicated that a large portion of periphyton is removed by the activity of the grazer rather than by direct ingestion. These results could be utilized to predict the impact of grazing on periphyton biomass.A contribution of the Groupe d'Écologie des eaux douces, Université de Montréal     

META‐ANALYSIS OF GRAZER CONTROL OF PERIPHYTON BIOMASS ACROSS AQUATIC ECOSYSTEMS1

Grazer control of periphyton biomass has been addressed in numerous experimental studies in all kinds of aquatic habitats. In this meta‐analysis, the results of 865 experiments are quantitatively synthesized in order to address the following questions: (i) Do lotic, lentic, and marine ecosystems differ in their degree of grazer control of periphyton biomass? (ii) Which environmental variables affect the degree of grazer control? (iii) How much does the result of these experiments depend on facets of experimental design? Across all ecosystems, the grazers removed on average 59% of the periphyton biomass, with grazing being significantly stronger for laboratory (65%) than for field (56%) experiments. Neither field nor lab experiments showed a significant difference among lotic, lentic, and coastal habitats. Among different taxonomic consumer groups, crustaceans (amphipods and isopods) and trichopteran larvae removed the highest proportion of periphyton biomass. Grazer effects increased with increasing algal biomass, with decreasing resource availability and with increasing temperature, especially in field experiments. Grazer effects also increased with increasing total grazer biomass in field experiments but showed the opposite trend in lab experiments, indicating a tendency toward overcrowded lab experiments. Other aspects of experimental design, such as cage type, size, and duration of the study, strongly affected the outcome of the experiments, suggesting that much care has to be placed on the choice of experimental design.     

Top‐down and bottom‐up control of periphyton by benthivorous fish and light supply in two streams

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纹沼螺对苦草生长的影响

在室外受控实验条件下,研究了不同密度(0,150,450个/m2)纹沼螺(Parafossarulus striatulus)对沉水植物苦草(Vallisneria spiralis)生长的影响。结果表明,螺类牧食活动促进了苦草生长,与无螺对照组相比,低密度螺处理组中苦草的相对生长率、株高均增加了20%;高密度组中苦草的相对生长率、叶片数和株高分别增加了28%、15%和27%。分析表明,纹沼螺通过牧食活动降低植物叶片上附着生物干重,从而促进了苦草生长。丰富了螺-草互利关系的研究内容,有助于加深理解水生态系统中生物之间的生态关系。     

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.     

Herbivory and intraspecific competition in a stream caddisfly population

Summary Field experiments were conducted to assess the effects of different population densities of the herbivorous caddisfly Helicopsyche borealis on periphyton biomass and on its own growth rate in Big Sulphur Creek, a third-order stream in northern California, USA. Stream enclosures were used to vary grazer density from one-eighth to twice natural density (1/8X–2X) in two experiments (35 d and 60 d), which spanned the period of most rapid larval growth. Periphyton biomass and chlorophyll a were inversely related to grazer density. Grazer densities of 1/8X–1/2X moderately reduced periphyton when compared to an ungrazed control, whereas densities of 1X–2X greatly depleted periphyton. The growth rate of H. borealis larvae declined with increasing larval density. Growth rate was highest at densities of 1/8X–1/2X; larvae grew more slowly at 1X and showed no growth at 2X. Weekly supplementation of periphyton to 1/4X, 1X, and 2X densities significantly increased the final body mass of larvae at 1X and 2X (by 26% and 50%, respectively, compared to unsupplemented larvae) but did not change the body mass of larvae at 1/4X. These results suggest that periphyton is a limiting resource in Big Sulphur Creek and that H. borealis larvae compete exploitatively for that resource. Intraspecific competition may be an important, but often overlooked, feature of many herbivore populations in streams.