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

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

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     

Striking convergence in the mouthpart evolution of stream-living algae grazers

This scanning-electron microscopic study demonstrates the convergent evolution of the mouthparts of various herbivorous stream animals (insects from different orders, an isopod, snails, fish, and a tadpole) feeding on epilithic algal pastures. This food source is rich but is often difficult to harvest. Nevertheless, a large number of species can live on it because they have evolved highly specialized mouthparts. There are four functional problems that an algae grazer has to solve: the algae must be removed from the stone, they have to be collected and crushed, and a current shield is needed to prevent the water flow sweeping away the food. Among the 30 algae grazers examined in this study, a limited number of morphological solutions have been found for each of these adaptational problems. There are multiple evolutionary pathways for mouthpart adaptation and even closely related species have often evolved different types of tools for the same function. This refects the existence of a certain amount of evolutionary scope. Such freedom of evolution is present, however, only at the beginning of the adaptiogenesis of an algae grazer. Once one of the evolutionary pathways is taken, further improvement of the mouthparts is possible only by the refinement of the ‘chosen’ type of tools. The consequence of this is that a large number of astonishing convergences have occurred in algae grazers that have independently trodden the same evolutionary pathway.     

Current velocity and habitat patchiness shape stream herbivore movement

Animal movements are influenced by the structure and arrangement of patches in a landscape. Most movement studies occur in terrestrial landscapes, though aquatic landscapes are equally heterogeneous and feature patches that differ in resistance to animal movements. Furthermore, the variable and highly directional flow of water over streambed landscapes is a unique environmental element, yet its constraint on animal movement is poorly understood. This study examines how habitat availability in a streambed landscape interacts with current velocity to affect movement patterns of two benthic grazers: glossosomatid caddisfly larvae (Agapetus boulderensis) and pulmonate snails (Physa sp.). Using experimental streambed landscapes, we found that Agapetus traveled farther as availability of smooth habitat (composed of low diatom turfs) increased compared to tall, structured filamentous stands, but only did so in slow current velocities. Swifter flows caused restricted movement of Agapetus and more upstream‐oriented paths, but only in smooth landscapes where the potential for flow refugia from filamentous stands was minimal. Similarly, increasing proportions of smooth habitat facilitated greater net displacement of Physa using more upstream‐oriented paths. Higher current velocities caused Physa to move faster, a pattern demonstrated only in smooth landscapes. Our results illustrate a strong interaction between benthic habitat structure and current velocity in shaping patterns of grazer movements in a streambed landscape. Our study also suggests that the flow of water be considered not only a strong environmental gradient in streams, but also an interactive landscape feature that can combine with streambed structure to determine the permeability of patches to the movement of benthic organisms. Landscape ecology has mainly focused on terrestrial environments, and this study offers insight into some of the unique processes that may shape animal movement in aquatic environments.     

Nitrogen processing by grazers in a headwater stream: riparian connections

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Macroinvertebrate grazers,current velocity,and bedload transport rate influence periphytic accrual in a field-scale experimental stream

Periphyton plays an important role in stream ecology, and can be sensitive to macroinvertebrate grazers, near-bed current velocity, and bedload abrasion. We manipulated conditions to examine influences on periphytic accrual in the St. Anthony Falls Laboratory Outdoor StreamLab in Minneapolis, MN, USA. Macroinvertebrate grazers were excluded from 27 of 65 clay tiles using electric pulses. We examined periphytic biomass accrual as a function of grazer presence, sampling run, and near-bed current velocity using ANCOVA. We found significant temporal differences between sampling runs but no significant effect of grazer presence. Along with a strong association between bedload transport rates and mean periphytic biomass, our results suggest that grazers are relatively unimportant in stream systems with high levels of physical disturbance from floods and associated sand bedload. However, the interaction between grazer presence and velocity was marginally significant. Regression analyses showed no relation between velocity and periphyton in the absence of grazers but a negative relation when grazers were present, suggesting that mechanical dislodgement of periphyton by grazers may increase with velocity. We conclude that grazers can have subtle effects on periphyton, particularly in streams with high bedload transport rates.     

Effects of removal of riparian vegetation on algae and heterotrophs in a Mediterranean stream

The effect of removal of a riparian strip on aquaticautotrophic (algae) and heterotrophic (bacteria,macroinvertebrates) organisms was monitored in aMediterranean stream during the canopy growing period.Community composition, biomass and metabolicactivities were compared with those recorded during apre-riparian removal period and in a forested stretchdownstream. Higher irradiance was associated with Cladophora increase in the logged section. Algalbiomass increased up to ten times, and productivitywas up to four times higher than in the pre-removalperiod and the forested section. Bacterialcommunities showed higher ectoenzymatic activities(-glucosidase, -xylosidase) in thelogged section than in forested conditions. Moreoverthe coincidence between the maxima of-glucosidase and chlorophyll-a suggeststhat bacterial activity was enhanced by the higheravailability of high-quality algal material. Themacroinvertebrate community had higher density andbiomass in the logged section than in the forestedsection and in the pre-removal period. Scrapers andfilterers become dominant after riparian removal,while shredders, predators and collectors did not showsignificant changes either between sites or periods.Responses of environmental variables and bioticcommunities indicate that the changes occurring in thestream due to riparian removal could be consideredbottom-up controlled, as increased illumination wasthe main mechanism responsible.     

Group size of feeding stream case-bearing caddisfly grazers and resource abundance

Several herbivorous insects are known to benefit from feeding in groups; however, little is known about (1) the resource conditions under which herbivorous insects perform group feeding and (2) the optimum population size to get any benefits by group feeding, for example, in terms of growth performance. To test the hypotheses that the benefits from group feeding change with resource level and population size, we performed field investigations and an enclosure experiment using the grazer caddisfly larva Micrasema quadriloba. The field investigations revealed aggregated distributions of larvae (indicator of aggregation, I_δ=4.1±1.55, aggregated density: 12.7±5.3 individuals per 3.1×3.1 cm² (mean±1 SD)) when periphyton was abundant on stream cobbles and random distributions (I_δ=1.0±0.11) when periphyton was scarce. In the enclosure experiment, the relative growth rate (RGR) of the larvae at each population size showed different tendencies at high and low periphyton abundance levels; RGR with abundant periphyton had a convex curve with a peak at intermediate population size, whereas RGR with scarce periphyton decreased linearly with increasing population size. The benefits from group feeding thus changed with resource level; larvae obtained high growth performance by group feeding behavior only when the resource was sufficiently abundant. The present study revealed not only that the optimum group size of larvae increased their growth performance, but also that this optimum group size occurred frequently in the field. We also discuss the mechanisms and benefits of group feeding by case-bearing caddisfly grazers.     

Relationship between littoral grazers and metaphytic algae in five softwater lakes

The zoobenthos communities within the littoral zones of five low alkalinity, Ontario lakes were examined during summer 1987. Detached algal clouds or metaphyton, dominated by algae from the family Zygnemataceae, accumulated over the summer in three of these lakes. Previous researchers have hypothesized that diminished grazing pressure due to extirpation of acid sensitive herbivores might enhance the development of metaphytic algae at pHs less than 6. No overt consistent differences existed among the abundances of small (cladocerans, cyclopoids, ostracods, oligochaetes, dipterans) or medium (amphipods, gastropods, emphemeropterans, trichopterans) grazers in study lakes with or without metaphyton. Notable differences did exist, however, with respect to what we term large grazers. Crayfish, algivorous fish (mostly cyprinids), and to a lesser extent tadpoles, were absent or rare in the three lakes with metaphytic algae but abundant in those without. If a reduction in grazing pressure can be implicated as a factor in enhanced metaphyton development, our correlative results suggest that it is the large grazers, particularly crayfish, that are the most likely candidates.     

Nutrients, algae and grazers in some British streams of contrasting pH

1. The relationship between algal biomass accumulation, invertebrate colonization, and stream-water pH was investigated in seven streams in three regions of England and Wales. Possible nutrient limitation of algal production at all sites was examined with diffusion substrata. 2. Periphyton assemblages on experimental substrata after 30 days were dominated by diatoms, notably Eunotia spp., at all sites. Algal pigment concentration (chlorophyll a and phaeopigments) was not correlated with stream-water pH, and mean concentrations on control (unenriched) substrata ranged from 0.08 to 1.94 μg cm^?2. 3. The growth response of periphyton to nutrient additions was site specific. Algal production was stimulated by nutrient additions at sites in the English Lake District and Llyn Brianne (south-west Wales), but not in the Ashdown Forest (southern England). 4. Larval Chironomidae were the main invertebrates retrieved from substrata at all sites. Within all three regions, larval abundance was positively related to algal pigment concentration (biomass). Abundance of the stonefly Nemurella pictetii was also positively correlated with algal biomass at the one site where it occurred. 5. Our results indicate that epilithic algal production in small, oligotrophic streams is unlikely to be determined primarily by pH. Neither do they support the view that an absence of grazers from acid streams is necessarily due to an inadequate food supply.     

The influence of crevice size on the protection of epilithic algae from grazers

1. This study investigated how the size of crevices might affect their effectiveness as refuges for diatom‐dominated algal assemblages from the grazing minnow Campostoma anomalum. 2. Crevice size was manipulated by making fired‐clay substrates, using moulds to produce eight substrates with pits from 1.17 to 22.0 mm diameter. Non‐pitted clay‐stones and limestone were also tested. Cages were used to control the access of Campostoma to arrays of the 10 different algal‐colonised substrates. The grazing treatments were: open and grazed, caged and ungrazed, and a grazed cage control. The experiment was replicated in eight large outdoor tanks. After 3 weeks, substrates were brushed and chlorophyll a concentrations of the removed algae and the algae remaining in pits were measured. 3. The experiment was field‐validated by exposing arrays of substrates to grazing Campostoma in five pools of a limestone stream. 4. The clay‐stone and limestone substrates accrued similar algal biomass and assemblages. 5. Smaller crevices provided more protection against grazing than larger crevices. Specifically, pits with openings smaller than 2 mm protected the enclosed algal assemblages in both the tank and field experiments. Larger pits provided less protection and pits over 7 mm in diameter were heavily grazed and may even be preferentially grazed by Campostoma. 6. None of the tested pit sizes were protective against larval chironomid grazers in the tank experiment, demonstrating that differences in the grazer size influence the effectiveness of crevice refuges.     

Colour and AOX removal from pulping effluents by algae

A mixed culture of algae was used to treat pulping mill effluent in terms of removing both colour and adsorbably organic halides (AOX). The removal of AOX from pulping effluent increased with increasing initial colour value of the effluent. However, for the total mill effluent (composed of both pulping and bleaching effluents), AOX removal was found to be independent of initial colour value, and was around 70%. Up to 80% removal of colour from pulping effluent was achieved within 30 days under continuous lighting conditions. It was found that algae reduced the colour of pulping effluent of relatively low initial colour more efficiently than that of high initial colour. Under simulated field lighting conditions, up to 60% colour removal from pulping effluent was observed after 60 days of exposure, whereas for the total mill effluent it was up to 64% after 45 days. Total organic carbon and lignin (UVA₂₈₀) were also removed to a significant extent, suggesting that the mechanism of colour removal might not be transformation of the coloured lignin molecules to non-coloured ones. Analysis of alkaline extraction of the algal biomass and material balance findings indicated that the main colour removal mechanism was metabolism rather than adsorption. The experimental results were also analysed using multiple regression techniques and a mathematical model was developed to express the removal of colour from pulping effluents in terms of initial colour value, exposure time and lighting periods as well as interactions between these variables. Received: 12 January 1999 / Revision received: 25 March 1999 / Accepted: 26 March 1999     

Potential functional redundancy and resource facilitation between tadpoles and insect grazers in tropical headwater streams

1. We quantified production and consumption of stream‐dwelling tadpoles and insect grazers in a headwater stream in the Panamanian uplands for 2 years to assess their effects on basal resources and energy fluxes. At the onset of our study, this region had healthy, diverse amphibian populations, but a catastrophic disease‐driven decline began in September 2004, which greatly reduced amphibian populations. 2. Insect grazer production was 348 mg ash‐free dry mass (AFDM) m^?2 year^?1 during the first year of the study and increased slightly to 402 mg AFDM m^?2 year^?1 during the second year. 3. Prior to amphibian declines, resource consumption by grazers (tadpoles and insects) was estimated at 2.9 g AFDM m^?2 year^?1 of algal primary production, which was nearly twice the estimated amount available. Insect grazers alone accounted for c. 81% of total primary consumption. During the initial stages of the declines, consumption remained at c. 2.9 g AFDM m^?2 year^?1, but only 35% of the available resource was being consumed and insect grazers accounted for c. 94% of total consumption. 4. Production and resource consumption of some insect grazers increased during the second year, as tadpoles declined, indicating a potential for functional redundancy in this system. However, other insect grazer taxa declined or did not respond to tadpole losses, suggesting a potential for facilitation between tadpoles and some insects; differential responses among taxa resulted in the lack of a response by insect grazers as a whole. 5. Our results suggest that before massive population declines, tadpoles exerted strong top‐down control on algal production and interacted in a variety of ways with other primary consumers. 6. As amphibian populations continue to decline around the globe, changes in the structure and function of freshwater habitats should be expected. Although our study was focused on tropical headwater streams, our results suggest that these losses of consumer diversity could influence other aquatic systems as well and may even reach to adjacent terrestrial environments.     

Nutrient removal from secondary effluent by filamentous algae

The nutrient removal potential of filamentous Oscillatoria sp. was quantitatively studied. Oscillatoria sp. showed satisfactory growth on secondary activated sludge effluent supplemented with 1% NaHCO₃. Continuous open culture of Oscillatoria sp. was kept stable using a continuous stirred tank equipped with a filter-separator. Kinetic constants Y_N and Y_P were 10.6 g cells/g NO₃-N and 37.7 g cells/g PO₄³⁻ respectively through the analysis of the results of continuous culture experiments. Monoalgal continuous culture of more than 90% purity could be maintained for 6 months without contamination. The harvested Oscillatoria cells were proven to have an excellent filterability. They also have excellent autoflotation. The amino acid composition of the Oscillatoria algal protein compares favorably with the tentative standard for ideal protein defined by FAO/WHO.     

Mercury removal by immobilized algae in batch culture systems

The accumulation and volatilization of mercury by non-immobilized and immobilizedChlorella emersonii have been studied in batch culture systems. Reduction in the mercury concentration in the growth medium by non-immobilized cells was highly dependent on inoculum density, whilst reduction in mercury concentration by immobilized cells was rapid at all inoculum densities. Mercury accumulation by immobilized cell biomass was significantly greater than by non-immobilized cells with 10⁶ and 10⁵ cells bead^–1 or ml^–1. Volatilization of mercury by non-immobilized cell systems was greatest at higher inoculum densities, whereas more mercury was volatilized from immobilized cell systems at lower inoculum densities, and was greatest with unstocked alginate beads. Thus, in immobilized systems, mercury removal from solution is complex and involves mercury accumulation by the cells and volatilization by the matrix and cells. Further studies of mercury accumulation and volatilization by unstocked immobilization matrices revealed that agarose volatilized much less mercury than alginate or agar. The precise mechanism of mercury volatilization by alginate remains unclear, though it is thought to be a chemical effect.     

Effect of pH on growth of acid stream algae

Five species of algae (Chlamydomonas applanata var. acidophila, Euglena mutabilis, Gloeochrysis turfosa, Hormidium rivulare, Stichococcus bacillaris) were isolated from a stream at pH 2·6–3·1, and their laboratory growth studied. Growth of all species could occur at pH values lower than those from which they were isolated, the lowermost limits being quite similar to those recorded for the particular species growing anywhere in England. Morphological differences were apparent with all five species at the lowermost pH values. These took place with Stichococcus bacillaris at pH values at which there was little reduction in growth rate, but with the other species obvious differences in morphology were correlated with a marked reduction in growth rate below the optimum rate. At the uppermost pH value tested, however, no obvious morphological differences were apparent. The effect of including 10% stream water in the medium was rather similar for all five species. No influence on growth rate was detectable at the lower pH values, but higher pH values led to a decrease in growth rate as compared with that found in medium lacking stream water.     

Ingestion and digestion of epilithic algae by larval insects in a heavily grazed montane stream

1. Epilithic algae grown on elevated or non-elevated ceramic tiles were exposed (to produce assemblages with different grazing histories) in a heavily grazed, montane stream in New Mexico, U.S.A. to Ameletus nymphs (Ephemeroptera) and Ecclisomyia larvae (Trichoptera) and the algal composition in insect faeces was compared to that on the tiles. Differences in grazing and digestion efficiency between grazers were then assessed and also differences in susceptibility to ingestion and digestibility among common algae. 2. Ordination of tile and faecal samples, using the relative abundance of common algae, revealed that: (i) algal assemblages on elevated vs. non-elevated tiles differed only slightly; (ii) the taxonomic composition of algae in faeces of both caddis and mayflies differed substantially from that on the tiles, indicating low grazing efficiency for some algal taxa; and (iii) the algal composition of faeces produced by caddis larvae and mayflies was similar, indicating little difference in grazing efficiency between them. However, some algal taxa were more susceptible to ingestion by caddisfly larvae when occurring on elevated tiles than on non-elevated tiles, suggesting that previous exposure to caddis grazing influenced assemblage attributes. 3. Although Ameletus and Ecclisomyia differed little in grazing efficiency, the percentage of diatoms that were dead after passage through the gut was greatest in the mayfly treatment, suggesting that mayflies digested diatoms more efficiently than the caddis. Analyses of differences in the condition of chloroplasts within diatoms in tile and faecal samples showed that losses of ’live‘ diatom cells (i.e. those containing full chloroplasts) during gut passage through mayflies equalled the increase, in faeces, of ’dead‘ (empty frustules) cells of all common diatoms. In contrast, some diatoms were digested inefficiently by caddis larvae. 4. Algae on elevated tiles contained a higher proportion of dead diatoms than those on non-elevated tiles, possibly because mayflies visited raised tiles more often and, consequently, ingested and defaecated cells at a higher rate in the absence of caddis larvae. Moreover, diatom taxa differed in the percentage of cells that were dead within tile assemblages, with populations of typically grazer-resistant taxa (e.g. Achnanthidium minutissimum, Planothidium lanceolatum and Cocconeis placentula var. euglypta) containing significantly more dead cells than grazer-susceptible taxa [e.g. small, chain-forming Fragilaria (= Staurosirella)]. This result suggests that a trade-off exists between ingestion vs. digestion resistance of microalgae. Both the ingestion and digestion efficiency of algivorous macroinvertebrates could influence the structure and function of algal assemblages. In heavily grazed systems, where algal cells are probably processed through grazer guts repeatedly, differential resistance to digestion among algae may be particularly important.     

Local effects of a sedentary grazer on stream algae

1. Larvae of the sedentary aquatic caterpillar Petrophila confusalis (Walker) construct silken retreats around which they feed; outside these clearly demarcated grazed areas, stream algae are exposed to a variety of mobile grazers. Comparisons of the algal community inside and just outside grazed zones were made for third- and fifth-instar Petrophila in the South Fork of the Eel River, California. 2. Densities of both the filamentous macroalga Cladophora and diatom assemblages were significantly reduced within the grazed areas during both larval instars. Grazing of diatoms was taxonomically non-selective. 3. In spring, the grazed zones were relatively large (mean = 22.7 cm²) and visibly increased epilithic spatial patchiness. 4. Per cent composition of diatom assemblages inside and outside the grazed areas differed during the third instar but not during the fifth instar. During the third instar, the grazed zone contained more Synedra ulna (a common immigrant) and less Gomphonema olivaceum (a late-successional species).     

Bacteria and algae in stream periphyton along a nutrient gradient

1. Stream riffles in southern Ontario and western Quèbec were sampled for biomass (58 stations from 51 streams) and production (22 stations from 21 streams) of algae and bacteria in periphyton to test the hypothesis that bacteria in benthic biofilms compete with algae for nutrients. 2. Algal and bacterial biomass were positively correlated, as were algal and bacterial production. Bacterial production was also positively correlated to algal and bacterial biomass, but the relationship was not significant. The ratio of algal to bacterial biomass did not vary with nutrients whereas algal production tended to increase with nutrients more rapidly than bacterial production. 3. Instream nitrogen concentrations explained 38–58% of the variability in algal biomass and production. Bacterial abundance explained an additional 9–29% of the residual variance in algal production and biomass. However, the relationship between bacterial abundance and algal production and biomass, once nutrients were taken into account, was positive, in contrast to the predicted effect of competition. 4. Hence, we reject our original hypothesis that bacteria in biofilms compete with algae for nutrients and instead suggest that bacteria and algae in biofilms coexist in an association that offers space and resources to sustain production of both groups of organisms.     

Distribution of benthic algae and macroinvertebrates along a thermal stream gradient

The distribution and abundance of benthic algae and macroinvertebrates were examined along a natural thermal gradient formed by hot springs in Little Geysers Creek, Sonoma Co., California, USA. Maximum water temperatures ranged from 52 °C at the uppermost station to 23 °C at a station 400 m downstream. Benthic chlorophyll a decreased exponentially from 2.5 g m^–2 at 52 °C to less than 0.1 g m^–2 at 23 °C, a pattern of decline also exhibited by algal phaeophytin. Blue-green algae dominated at higher temperatures but were replaced by filamentous green algae and diatoms at lower temperatures.Macroinvertebrates were absent at temperatures 45 °C; the highest density (> 150 000 m^–2, mainly Chironomidae) occurred at 34 °C, whereas biomass was highest (4.6 g m^–2, as dry weight) at 23 °C and species richness (15 species) was highest at 27 °C. The two predominant macroinvertebrate populations (the midge Tanytarsus sp. and the caddisfly Helicopsyche borealis) occurred at sites that were several degrees below their lethal thermal threshold, suggesting that a temperature buffer is maintained.