Predatory fish impact on competition between stream insect grazers: a consideration of behaviorally- and density-mediated effects on an apparent coexistence pattern

A manipulative field experiment and theoretical analyses of a simple competition model were used to show how exploitative competition between a caddisfly (Glossosoma sp.) and three mayfly grazers (Ameletus sp., Baetisthermicus and Cinygmula sp.) was mediated by a predatory fish, freshwater sculpin (Cottus nozawae). The field experiment followed a two‐factorial design, with Glossosoma densities (natural vs reduced) and sculpin presence (present vs absent) as treatments. Diet analysis revealed that all four prey species were eaten under the natural condition and the sculpin preferred mayfly grazers to Glossosoma. Our experiment showed that although mayfly densities in the presence of either sculpin or Glossosoma were lower than in the no‐sculpin plus reduced‐Glossosoma treatment, no difference in mayfly densities was found between the following three treatments: sculpin plus natural‐Glossosoma, no‐sculpin plus natural‐Glossosoma, and sculpin plus reduced‐Glossosoma. These results indicated that fish predator produced no effects additive to the competitive operation on the mayfly density, and also that competitive operation of Glossosoma on the mayfly densities produced no additional effects to fish predators. In addition, although the competitive effect of the mayflies on Glossosoma could not be manipulated in isolation, the density of Glossosoma in the presence of sculpin was greater than that in the absence of sculpin in the natural‐Glossosoma treatment. Thus, the densities of these competing grazers either stayed the same or increased in the presence of the fish predator relative to predator‐free treatments. A theoretical model, developed to explain the experimental results here, predicted that the densities of the two competing prey under predation pressure could be simultaneously greater than those under predator absent conditions when the behaviorally‐mediated effects of the predator were strongly operative. Although we were unable to distinguish experimentally the two different effects of predator on the prey competition, the behaviorally‐ and density‐mediated effects, the reality of the behaviorally‐mediated effects in the experiment was discussed.     

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.     

The invasive New Zealand mudsnail, <Emphasis Type="Italic">Potamopyrgus antipodarum</Emphasis>, is an effective grazer of algae and altered the assemblage of diatoms more than native grazers

Exploitation of shared resources often mediates the impacts of invasive species on native species. In a field experiment, we compared the ability to graze periphyton and the genera of diatoms removed by the invasive New Zealand mudsnail, Potamopyrgus antipodarum, a native caddisfly larva (Brachycentrus sp.), and a native mayfly nymph (Ephemerella sp.) over 1 week. P. antipodarum removed as much or slightly more periphyton than the native grazers, depending on whether chlorophyll a or ash-free dry mass was used to measure periphyton biomass. When we examined the diatoms in the periphyton, P. antipodarum altered the diatom assemblage more than the native grazers. Effective grazing of periphyton by P. antipodarum may impact native grazers by consuming shared algal resources. In particular, because Ephemerella sp. were also effective grazers, these mayflies may compete for periphyton with P. antipodarum in the western United States. Taken together, these results suggest that ability to procure food resources may contribute to the invasion success of P. antipodarum.     

Alterations in the grazing activities of cased caddisfly larvae in response to variations in predation risk and resource level

Flexible behavioral response to avoid encountering predators has never been reported in lotic, cased caddisfly larvae with low mobility. However, an earlier laboratory experiment found the growth of such a caddisfly species, Glossosoma sp., decreased in the presence of a predatory sculpin, Cottus nozawae. We conducted laboratory experiments to test whether Glossosoma responses to sculpin varied according to resource level and/or time of day. Lower periphyton biomass resulted in increased movement at any time of day, but was followed by decreased grazing success of Glossosoma in the morning when they were sampled. Although sculpins were active in the morning, evening and night but never in the afternoon, movements of Glossosoma were suppressed by sculpin stimuli only in the morning, when food intake of Glossosoma also decreased, regardless of periphyton biomass. Glossosoma reduced the risk of predation by sculpin by flexibly controlling its grazing activities during the most risky period of the day (i.e. morning). Even in the morning in the presence of sculpin, lower resource levels resulted in longer movement distances similar to those in predator-free conditions, suggesting that lower resource availability promotes predation risk in natural streams inhabited by predatory sculpin.     

Does light intensity modify the effect mayfly grazers have on periphyton?

1. A factorial experiment was conducted in artificial outdoor streams to quantify the effects of irradiance (two levels) and two mayfly grazers (four densities of each) on periphytic community structure. The mayflies were Ecdyonurus venosus (Heptageniidae), a grazer using brushing mouthparts, and Baetis spp. (Baetidae) a grazer which uses mandibles and maxilla to scrape and gather periphyton. The experiment ran for 16 days. 2. Grazer densities in channels approximated those existing in a shoreline habitat in the River Sihl, Switzerland. Light treatments were natural (daily mean = 810 μmol m^–2 s^–1) and shaded (daily mean = 286 μmol m^–2 s^–1). 3. Higher irradiance increased total algal abundance by a factor of 4. Algae most affected were prostrate/motile and erect diatoms, filamentous chlorophytes and Hydrurus foetidus. 4. Both species of mayfly reduced periphytic and algal biomass. Mayfly–mayfly interactions, however, were associated with statistical increases in algal biovolume and chlorophyll-a content, indicating that the two grazers may have interfered with one another as their densities increased. The mayfly–mayfly interaction did not influence periphytic ash-free dry mass (AFDM). Light modified the influence of Ecdyonurus such that this mayfly produced greater reductions in algal biovolume under high irradiance. 5. Despite efforts to exclude other grazers, chironomids colonized experimental channels. Chironomid biomass was approximately eight times less than mayflies across treatments and was positively correlated with all measures of periphytic abundance, suggesting that these grazers were responding to periphyton rather than controlling it. Chironomids were also associated with an increase in the abundance of diatoms having a prostrate/motile physiognomy. The only physiognomy to show a negative relationship with chironomid biomass was the thallus type, a form which comprised less than 1% of the algal biovolume across channels. 6. Ecdyonurus and Baetis had distinct influences on algal physiognomy. Ecdyonurus, for example, reduced adnate, stalked and Achnanthes-type physiognomies, but was associated with a significant increase in the abundance of filamentous chlorophytes (primarily Ulothrix sp.). Baetis reduced erect, Achnanthes-type and thallus physiognomies. Neither mayfly influenced the abundance of prostrate/motile diatoms; a physiognomy that comprised 21% of the algae in channels. 7. Light and mayfly interactions affected algal community structure. The interaction of Ecdyonurus with light had a negative effect on erect diatoms, filamentous chlorophytes and the thallus physiognomy, but a positive effect on stalked and Achnanthes-type physiognomies. Baetis interacting with light had a positive effect on adnate diatoms. 8. Although both mayfly taxa influenced periphytic community structure, physiognomy was not a good predictor of algal susceptibility to grazing. The type of substratum to which an alga is attached (detritus or algal filaments vs hard surfaces) and location within the periphytic matrix may be better indicators of vulnerability to grazing than physiognomy.     

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.     

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)     

Protozoan communities in chalk streams

This study assessed the individual effects of three mayflies (Paraleptophlebia sp., Ephemerella subvaria McDunnough and Epeorus sp.) and one caddisfly (Psilotreta sp.) on periphyton communities associated with clay tiles and leaves. Algal densities were estimated for leaf discs and tiles from experimental chambers (with individual grazers) and control chambers (i.e., no grazers). Scanning electron micrographs (SEM) of leaf discs and tiles also were taken for all mayfly grazing experiments. Densities of algae on leaf discs were two to five times lower than on tiles. Mouthpart morphology influenced how different insects grazed the periphyton community. Paraleptophlebia had typical collector-gatherer mouthparts and had no effect on diatom densities associated with leaves whereas diatom densities on grazed tiles were higher than densities on tiles from control chambers. Epeorus had brusher mouthparts and had little impact on diatom densities regardless of substratum type. The other two grazers had the blade-like mandibles of a scraper. Psilotreta did not reduce the numerical abundance of diatoms on either substratum, but did alter community structure by significantly reducing densities of stalked Gomphonema olivaceum and large species of Navicula and Nitzschia; densities of smaller diatoms (Achnanthes spp) increased. However, E. subvaria reduced densities of most algal species regardless of size on both substrata and also significantly altered community structure. SEMs of substrata grazed by mayflies showed reductions in fungal hyphae on all grazed leaf discs, decreases in filamentous algal forms on grazed tiles, and greatly shortened stalks of G. olivaceum (Paraleptophlebia only). Thus, periphyton communities are different on leaves versus tiles and grazers with different mouthpart morphologies have varying effects on both algal and heterotrophic microbial community structure.     

Removal of settled sediments and periphyton from macrophytes by grazing invertebrates in the littoral zone of a large oligotrophic lake

• 1 The resistance and resilience of littoral zone communities to sedimentation will depend both on the extent to which sediment deposition affects productivity, and on interactions within the communities. A series of hypotheses were set up and tested to examine interactions and feedback mechanisms among deposited sediments, periphyton, macrophytes and grazers in a large oligotrophic lake subject to fluctuating sediment loadings.
• 2 Although sediments incorporated into periphyton reduced light availability to macrophytes, periphytic algae were generally the dominant light absorbing component under natural conditions. When grazers were absent, both sediments incorporated in the periphyton and periphytic algal densities increased, and both were then important in reducing light available to macrophytes.
• 3 Grazing rate and assimilation efficiency for the dominant grazer, the prosobranch gastropod Potamopyrgus antipodarum, increased with increasing sediment content under natural lake conditions to reach a maximum at 10 mg sediment cm^?2.
• 4 An increase in sediment incorporation into periphyton films resulted in an increased grazing rate and hence grooming of sediments from macrophytes.
• 5 Grazing invertebrates can play a major role in maintenance of littoral communities by continuously grooming macrophyte hosts of periphytic algae and settled sediments.

     

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.     

The relationship between food density and short term assimilation rates in Potamopyrgus antipodarum and Deleatidiumsp.

Much of the variation in individual growth rates can be attributed to differences in individual feeding rates. Therefore, in order to build predictive models of individual, or population growth, the factors influencing an individual's feeding rate must be described. An important determinant of the feeding rate is the relationship between the local abundance of food and the individual's ingestion rate – otherwise known as the functional response. We determined functional responses for two species of invertebrate grazers: the snail Potamopyrgus antipodarum and the mayfly Deleatidium sp., by measuring their assimilation rate with increasing densities of radiolabelled periphyton. The assimilation rates were consistent with the Holling Type II or Michelis Menten functional response curve. The parameters of the functional response yielded estimates of the search area and handling time for the stream invertebrates. Our functional response data indicate that the half-saturation food density for P. antipodarum and Deleatidium sp. were 980 mg and 3200 mg AFDM m^–2, respectively, suggesting that Deleatidium growth may be subject to food limitation more often than is P. antipodarum – despite the lower assimilation efficiency of the latter species.     

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.     

The influence of periphyton biomass and density on grazing in Physella virgata

We studied how differences in periphyton colonization interval and snail density affected grazing rates in Physella virgata, and whether snails controlled periphyton biomass. Both egestion rates and incorporation rates of ¹⁴C labeled periphyton were estimated in laboratory experiments. Periphyton biomass increased with field colonization interval in all experiments, but did not consistently influence estimates of grazing rate. However, increased periphyton abundance in one of the experiments could still explain higher grazer rates in that year, although larger snail body size is a confounding explanation. Increased snail density also resulted in decreased grazing rates, as observed in earlier studies with this snail species, as well as in studies with other snail grazers. Our results suggest grazing rates and resulting impacts may change seasonally with variation in either periphyton biomass, grazer life-history stage or population density.     

Independent and interactive effects of nutrients and grazers on benthic algal community structure

Algal responses to nutrients, grazing by Helicopsyche borealis, and concurrent grazing by Helicopsyche and Baetis tricaudatus were examined in recirculating stream chambers. Alagl communities, dominated by Achnanthes minutissima, Cocconeis placentula, and Synedra ulna, were primarily phosphorus-limited. Algal populations responded after only 6 days of nutrient enrichment. Initially, both the adnate diatom Cocconeis and erect diatom Synedra showed positive response to nutrient enrichment. Accumulation of algal biomass between day 3 and 6 in the P enriched treatment was resulted primarily from the growth of Synedra, an overstory rosette-like diatom colony. Such a shift in dominant growth from adnate to erect diatoms is a general phenomenon in periphyton succession in the absence of disturbance. Algal species showed differential responses to an increase of Helicopsyche densities. The accrual rate of Achnanthes continuously decreased with increasing grazer densities. The accrual rates of both Cocconeis and Synedra declined but reached plateaus between medium and high grazing densities. Baetis effectively and exclusively depressed Synedra and had no significant impact on Cocconeis. After concurrent grazing, algal communities were mainly dominated by Cocconeis (approximately 80% of total algal biovolume). The grazer' s mouth structures, grazing efficiencies, and mobility may account for the differential effects of concurrent grazing on algal communities. Significant interactive effects of P and grazing by Helicopsyche indicated that both nutrient addition and grazing may exert significant impact on algal communities. However, grazing may have a much stronger effect on algae than nutrients. Our results indicate that enhancement of algal biomass by P was dampened by grazing activities and that P had no effect on algal biomass in the presence of grazers.     

Comparative performance of benthic diatom indices used to assess river water quality

Grazer-periphyton interactions were investigated in 11 laboratory streams holding a range of densities of three herbivore taxa during a 32-d experiment. Effects of grazers on algae were strongest with Dicosmoecus gilvipes caddisflies, intermediate with Juga silicula snails, and weakest with Baetis spp. mayflies. Algal standing crop, export, and gross primary production declined logarithmically with increasing grazer density. Algal turnover rate, however, increased with grazer abundance. At high densities of all grazers, responses in most algal parameters converged, suggesting that high grazing pressure, regardless of taxon, will similarly affect periphyton. Growth of both Dicosmoecus caddisflies and Juga snails was density-dependent, with the highest growth rates occurring at the lowest densities. Caddisflies displayed high growth rates but low efficiency in resource use. Snails had lower growth rates but were more efficient in resource use. The coexistence of Dicosmoecus and Juga, or other competing herbivores, in natural streams may be related to these fundamental differences in life history strategies.Department of Fisheries and Wildlife     

Fish foraging effects on benthic assemblages along a warm-temperate stream: differences among drift feeders, benthic predators and grazers

In streams, physical and biotic conditions change from the headwaters to the mouth, shaping longitudinal patterns in community structure. We examined how fish foraging effects on periphyton and benthic invertebrates changed along a longitudinal gradient of a warm-temperate stream in southwestern Japan. We established three study sites according to changes in the composition of fish feeding guilds (upper site characterized by drifting-invertebrate feeders, Oncorhynchus masou ; middle site by benthic invertebrate feeders, Rhinogobius spp.; lower site by the presence of periphyton grazers, Sicyopterus japonicus ), and performed two manipulative experiments to examine effects of different fish assemblages on periphyton and benthic invertebrate abundances. Results of an exclosure experiment suggested that fishes had no effect on the benthic assemblages at the upper and middle sites whereas fishes reduced the abundances of both periphyton and invertebrates on stone surfaces at the lower site, where both benthic invertebrate feeders and grazers inhabited. A subsequent enclosure experiment showed that the reduction of invertebrate densities at the lower site was caused by the grazers rather than benthic invertebrate feeders. These experimental results suggested that effects of fishes on benthic assemblages are intensified downstream, owing to the occurrence of the grazing fish. Furthermore, observational data based on field sampling suggested that such grazing effects were reflected in longitudinal patterns in periphyton and invertebrate abundances. Overall results emphasize an important role of the grazing fish ( S. japonicus ) in shaping longitudinal patterns in benthic assemblage structure.     

Periphyton density and shading in relation to tidal depth and fiddler crab activity in intertidal seagrass beds of the Banc d'Arguin (Mauritania)

Periphyton development was studied on microscopic glass slides and leaves of Zostera noltii Hornem. in an intertidal area in the Banc d'Arguin (Mauritania). The effects of shading, tidal depth and grazing activities by the fiddler crab Uca tangeri Eydoux were evaluated. For all experiments, periphyton ash content was high (52–93%) and ash-free dry weight ranged between 0.10–0.63 mg cm^–2. Slides accumulated more periphyton than leaves.Artificial shading (62–99%) for 13 days had no effect on periphyton densities on leaves. Increased tidal depth resulted in higher ash-free dry weight on slides, but in lower ash-free dry weight on leaves. Significant variation along the coastline also existed. The effect of fiddler crabs was studied using exclosures. Presence of fiddler crabs reduced periphyton density on slides, whereas light transmittance was increased. On leaves, no significant fiddler crab effect was found. This difference between leaves and slides was probably caused by a storm at the day before the end of the experiment, and by the higher periphyton density on slides as compared with leaves. As visual inspection during the experiment showed clear differences in appearance of leaves inside and outside the exclosures, the storm probably sloughed off mainly the older leaves, i.e. those on which the differences in periphyton cover were the highest.It is hypothesized that periphyton accumulation is higher with increased tidal depth, whereas fiddler crab grazing pressure also increases in this direction. The result is a decreased periphyton density with increased tidal depth.The presently found light extinction coefficients (mean 0.8 m^–1) and periphyton light attenuance (up to 25%) in Banc d'Arguin are not likely to affect seagrass leaf growth.     

Indirect effects of fish on macrophytes in Bays Mountain Lake: evidence for a littoral trophic cascade

Summary We began this experiment to test specific hypotheses regarding direct and indirect effects of fish predation on the littoral macroinvertebrate community of Bays Mountain Lake, Tennessee. We used 24 m² enclosures in which we manipulated the presence and absence of large redear sunfish (Lepomis microlophus>150 mm SL), and small sunfish (L. macrochirus and L. microlophus <50 mm SL) over a 16-mo period. Here we report on effects of fish predation on gastropod grazers that appear to cascade to periphyton and macrophytes.Both large redear sunfish and small sunfish maintained low snail biomass, but snails in fish-free controls increased significantly during the first 2-mo of the experiment. By late summer of the first year of the experiment, the difference in biomass between enclosures with and without fish had increased dramatically (>10×). Midway through the second summer of the experiment, we noted apparent differences in the abundance of periphyton between enclosures containing fish and those that did not. We also noted differences in the macrophyte distribution among enclosures. To document these responses, we estimated periphyton cover, biovolume and cell size frequencies as well as macrophyte distributions among enclosures at the end of the experiment. When fish were absent, periphyton percent cover was significantly reduced compared to when fish were present. Periphyton cell-size distributions in enclosures without fish were skewed toward small cells (only 12% were greater than 200 m³), which is consistent with intense snail grazing. The macrophyte Najas flexilis had more than 60 x higher biomass in the fish-free enclosures than in enclosures containing fish; Potamogeton diversifolius was found only in fish-free enclosures. These results suggest a chain of strong interactions (i.e. from fish to snails to periphyton to macrophytes) that may be important in lake littoral systems. This contrasts sharply with earlier predictions based on cascading trophic interactions that propose that fish predation on snails would enhance macrophyte biomass.     

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.     

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.