Hydraulic influences on periphyton and benthic macroinvertebrates: simulating the effects of upstream bed roughness

1. Hydraulic conditions, periphyton biomass and invertebrate communities were compared on artificial substrates exposed to a range of upstream roughness conditions across an area of uniform current velocity and depth in a gravel-bedded river. The effect of river bed roughness was simulated by installing roughness elements upstream of artificial substrates.
2. Increasing upstream roughness reduced the average near-bed velocity above the substrates and increased short-term variability in velocity (i.e. turbulence).
3. Periphyton chlorophyll a density showed a general decline with near-bed velocity and was significantly lower on the substrates exposed to the river bed reference and 0 mm roughness treatments than the 110 mm roughness elements. Chlorophyll a was also negatively correlated with the abundance of larger collector-browsing invertebrates. This indicates that effects of the changes in hydraulic conditions on invertebrates may have contributed to the observed treatment effects on periphyton.
4. Invertebrate abundance and diversity declined with increasing upstream roughness. Filter-feeders, collector-browsers and predatory invertebrates all declined in abundance with increasing upstream roughness, but the effect was strongest for filter-feeders. Eight of the nine most common taxa showed significant treatment effects. The orthoclad chironomid, Eukiefferiella sp., was not influenced strongly by upstream roughness, but its abundance was correlated significantly with periphyton biomass.     

Hydraulic parameters and benthic invertebrate distributions in two gravel-bed New Zealand rivers

1. Benthic invertebrates were sampled over a matrix of about eighty combinations of mean velocity (10–150cms^?1) and depth (10–150cm) in two rivers that differed in substrate size variability. Vertical velocity profiles were measured at each sample site and substratum roughness was measured and estimated from percentage cover by stone size classes. The influence of depth on periphyton biomass was also measured. 2. The hydraulic and substrate data were used to investigate the correlations between conventional (mean velocity, depth, substrate size) and complex hydraulic variables (Froude number, shear velocity, and water column and boundary Reynolds number) that were either calculated from direct measurements or inferred from mean velocity, depth, kinematic viscosity and substrate roughness. The ecological relevance of these hydraulic variables was investigated by comparing their degree of correlation with invertebrate densities and community metrics. 3. The invertebrate variables had similar correlations with mean velocity and the complex near-bed hydraulic variables in the river with uniform cobble substrates. In the river with diverse substrates, however, average correlations with Froude number, and inferred shear velocity and boundary Reynolds number were 25–45% higher than with velocity. Of all the individual hydraulic parameters, the boundary Reynolds number, calculated from simple measures, was most strongly correlated with benthic invertebrate distributions and taxa richness. However, invertebrate distributions were more strongly correlated with predictions of multiple regression models, incorporating substrate size, depth and mean velocity, than with any single hydraulic variable. 4. Hydraulic influences on food availability and oxygen concentration in the benthos are likely mechanisms affecting the hydraulic preferences of several taxa. Lower periphyton biomass with depth, partly attributable to light attenuation, appeared to have a non-hydraulic influence on a collector-browser species.     

Effects of prey dispersal on predator–prey interactions in streams

1. We studied the effect of mesh size (6 and 3 mm) on interactions between brown trout ( Salmo trutta ) and benthic invertebrates in enclosures placed in a stream in southern Sweden. We also compared how different prey exchange rates affected interactions between trout and invertebrates.
2. Trout had strong impacts on some benthic taxa, and different mesh sizes produced different patterns. Trout affected the abundance of 10 of the 21 taxa examined, six in enclosures with 3 mm mesh and six in enclosures with 6 mm mesh. The abundance of nine of the prey taxa was lower in the presence of trout, only leptocerids were more numerous in the presence of trout.
3. Our measurements of prey immigration/emigration, together with trout diet data, suggest that direct consumption by trout, rather than avoidance behaviour by prey, explains most decreases in prey abundance. There was avoidance behaviour by only two of the twenty-one prey taxa, with trout inducing emigration of the mayflies Baetis rhodani and Paraleptophlebia sp.
4. Trout indirectly increased periphyton biomass in both 3 and 6 mm enclosures. The effect of trout on periphyton was probably due to strong effects of trout on the grazer, Baetis rhodani , Heptagenia sp. and Paralepthoplebia sp.
5. Our results suggest that mesh size, through its effects on exchange rates of prey, may affect interactions between predators and prey in running waters, but that the effects of dispersal and predation on invertebrates are taxon specific.     

Patterns in benthic food webs: a role for omnivorous crayfish?

1. The biomass and species richness of macrophytes and invertebrates in artificial ponds at two sites in southern Sweden (twenty-one ponds at each site) were investigated. Alkalinity was high at one site (H ponds) and low at the other site (L ponds). The ponds chosen had different densities of signal crayfish (Pacifastacus leniusculus), with mean crayfish abundance (estimated by trapping and expressed as catch per unit effort) significantly higher in the L ponds (10.7) than in the H ponds (4.9). Macrophytes, invertebrates, the amount of periphyton on stones and the organic content of the sediment were determined in each pond. 2. Macrophyte biomass, cover and species richness declined with increasing crayfish density. Macrophyte species composition differed between ponds and was related to crayfish abundance. 3. The total biomass of invertebrates and the biomass of herbivorous/detritivorous invertebrates declined with increasing crayfish abundance, but the biomass of predatory invertebrates declined only in the L ponds. The relative biomass of Gastropoda and Odonata declined in ponds where crayfish were abundant. In ponds where crayfish were abundant the invertebrate fauna was dominated by sediment-dwelling taxa (Sialis (H and L ponds) and Chironomidae (H ponds)). 4. The number of invertebrate taxa in macrophytes declined with increasing crayfish abundance. The percentage of macrophyte-associated invertebrate taxa differed between ponds, but also between sites. The relative biomass of Gastropoda declined in H ponds where crayfish were abundant. In H ponds Trichoptera or Gammarus sp. and Heteroptera dominated where crayfish were abundant, whereas Odonata dominated in L ponds with abundant crayfish. 5. The organic content of the sediment decreased in ponds with high crayfish densities, while the amount of periphyton on stones was not related to crayfish density. 6. We conclude that the signal crayfish may play an important role as a keystone consumer in pond ecosystems, but lower trophic levels did not respond to changes in the abundance of the crayfish according to the trophic cascade model. Omnivorous crayfish may decouple the cascading effect.     

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.     

Periphyton responses to invertebrate grazing and riparian canopy in three northern California coastal streams

SUMMARY. 1. Field experiments were conducted to examine the impact of grazing invertebrates on periphyton biomass in twenty-one pools across three northern California coastal streams (U.S.A.): Big Sulphur Creek, the Rice Fork of the Eel River, and Big Canyon Creek. Periphyton accrual on artificial substrate tiles was compared in each stream between two treatments: those elevated slightly above the stream bottom to reduce access by grazers (= platforms) and those placed directly on the stream bottom to allow access by grazers (=controls).
2. Crawling invertebrate grazers (cased caddisflies and snails) were numerically dominant in each stream (86% of all grazers in Big Sulphur Creek, 61% in the Rice Fork, 84% in Big Canyon Creek). Platforms effectively excluded crawling grazers, but were less effective in excluding swimming mayfly grazers (Baetidae).
3. Periphyton biomass (as AFDM) on tiles was significantly lower on controls compared to platforms for the Rice Fork, an open-canopy stream, and Big Sulphur Creek, a stream with a heterogeneous canopy. In contrast, no grazer impact was found for Big Canyon Creek, a densely shaded stream. Here, extremely low periphyton biomass occurred for both treatments throughout the 60 day study.
4. The influence of riparian canopy on periphyton growth (i.e. accrual on platforms), grazer impact on periphyton, and grazer abundance was examined for Big Sulphur Creek. As canopy increased (15–98% cover), periphyton biomass on platforms decreased. In contrast, canopy had little influence on periphyton accrual on controls; apparently, grazers could maintain low periphyton standing crops across the full range of canopy levels. The abundance of one grazer species, the caddisfly Gumaga nigricula , was highest in open, sunlit stream pools; abundance of two other prominent grazers, Helicopsyche borealis (Trichoptera) and Centroptilum convexum (Ephemeroptera), however, was unrelated to canopy.     

The influence of sediment mobility and channel geomorphology on periphyton abundance

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Periphyton dynamics in a floodprone prealpine river: evaluation of significant processes by modelling

1. Periphyton chlorophyll a was measured at weekly or 2 weekly intervals from October 1992 to March 1994 at four sites in a Swiss prealpine gravel bed river that was frequently disturbed by unpredictable spates.
2. To evaluate the dominant processes that control periphyton biomass, measured data were compared with a set of simulations from an empirical dynamic periphyton model. Different combinations of process hypotheses were systematically activated and deactivated in order to assess their importance.
3. The simplest model leading to an acceptable agreement with measured data employs a biomass-dependent growth rate, a detachment rate directly proportional to discharge and biomass, and a catastrophic loss rate during bed moving spates. Terms describing light or temperature dependence had a minor effect on the model fit.
4. The model describes the temporal pattern of the periphyton biomass as a series of growth curves periodically truncated by spates. Within the uncertainties of the measurements, mainly caused by the spatial heterogeneity of periphyton, the biomass recovered along deterministic trajectories.
5. Sensitivity analyses with respect to model parameters and model structure showed that site-specific model parameters could not be unequivocally determined, and that the model yields similar results with slightly different formulations of processes. This indicates that the data base with respect to periphyton biomass was too small for a unique identification of model details but that the main conclusions on the significance of processes did not depend on arbitrary choices of the model formulation.     

Periphyton dynamics in a floodprone prealpine river: evaluation of significant processes by modelling

1. Periphyton chlorophyll a was measured at weekly or 2 weekly intervals from October 1992 to March 1994 at four sites in a Swiss prealpine gravel bed river that was frequently disturbed by unpredictable spates.
2. To evaluate the dominant processes that control periphyton biomass, measured data were compared with a set of simulations from an empirical dynamic periphyton model. Different combinations of process hypotheses were systematically activated and deactivated in order to assess their importance.
3. The simplest model leading to an acceptable agreement with measured data employs a biomass-dependent growth rate, a detachment rate directly proportional to discharge and biomass, and a catastrophic loss rate during bed moving spates. Terms describing light or temperature dependence had a minor effect on the model fit.
4. The model describes the temporal pattern of the periphyton biomass as a series of growth curves periodically truncated by spates. Within the uncertainties of the measurements, mainly caused by the spatial heterogeneity of periphyton, the biomass recovered along deterministic trajectories.
5. Sensitivity analyses with respect to model parameters and model structure showed that site-specific model parameters could not be unequivocally determined, and that the model yields similar results with slightly different formulations of processes. This indicates that the data base with respect to periphyton biomass was too small for a unique identification of model details but that the main conclusions on the significance of processes did not depend on arbitrary choices of the model formulation.     

Effects of substratum stability on diversity of stream invertebrates during baseflow at two spatial scales

1. The effects of substratum stability on the diversity of stream invertebrates were assessed at two spatial scales in a Japanese stream during baseflow, from May to June 1998. Deposition and erosion were examined separately as distinct elements of substratum stability by a newly developed method using small steel pins. Stream invertebrates were sampled after 28 days of measurement of substratum stability. We also measured physical environmental variables, current velocity and depth, and food resource parameters including periphyton and particulate organic matter (POM) standing crops.
2. At the scale of the habitat patch, the effects of substratum stability on invertebrates were overwhelmed by those of POM standing crop. Moreover, higher taxon richness at high abundance may simply result from a higher likelihood of more taxa being included in samples. Therefore, this small scale revealed no role for substratum stability in explaining spatial pattern of community diversity.
3. At the reach scale ( n =10), taxon richness and evenness peaked at an intermediate level of deposition, whereas invertebrate abundance did not show any significant relationship. This result, and the pattern of relative abundance of common taxa, implies that the diversity of stream invertebrates was determined by subtle substratum movements and by the habitat preference of each taxon.
4. The difference in the determinant of community parameters between the habitat patch and the reach affirm the importance of a cross-scale analysis to choose an appropriate spatial scale for investigating the community structure of stream invertebrates. Prominent effects of substratum stability, particularly the deposition of substratum particles, during baseflow suggest that subtle and constant movement of small substratum particles can contribute to determine the diversity of stream invertebrates.     

The response of two submerged macrophytes and periphyton to elevated temperatures in the presence and absence of snails: a microcosm approach

Global warming may affect snail–periphyton–macrophyte relationships in lakes with implications also for water clarity. We conducted a 40-day aquaria experiment to elucidate the response of submerged macrophytes and periphyton on real and artificial plants to elevated temperatures (3°C) under eutrophic conditions, with and without snails present. With snails, the biomass and length of Vallisneria spinulosa leaves increased more at the high temperature, and at both temperatures growth was higher than in absence of snails. The biomass of periphyton on V. spinulosa as well as on artificial plants was higher at the highest temperature in the absence but not in the presence of snails. The biomass of Potamogeton crispus (in a decaying state) declined in all treatments and was not affected by temperature or snails. While total snail biomass did not differ between temperatures, lower abundance of adults (size >1 cm) was observed at the high temperatures. We conclude that the effect of elevated temperature on the snail–periphyton–macrophyte relationship in summer differs among macrophyte species in active growth or senescent species in subtropical lakes and that snails, when abundant, improve the chances of maintaining actively growing macrophytes under eutrophic conditions, and more so in a warmer future with potentially denser growth of periphyton.     

Initial colonization of periphyton on natural and artificial apices of Myriophyllum heterophyllum Michx

SUMMARY. 1. The initial colonization of periphyton on natural and artificial apices of Myriophyllum heterophyllum Michx was compared at three depths in the littoral zone of Lake Winnipesaukee, a soft-water New England lake. After a 1 week incubation period, the apices were sampled and the periphyton was removed and counted as numbers of periphyton organisms per centimetre of stem.
2. Initial colonization on both substrates was characterized by diatoms, particularly small single-celled species, throughout the spring and summer. Blue-green and green algae occurred in low numbers on both substrates during mid-summer, usually forming a greater percentage of the population of the natural apices. Blooms of the green alga Zygnema sp. dominated both substrates in late August.
3. Community composition generally did not differ significantly by depth or substrate on the natural and artificial apices: however, total abundance was significantly greater on the natural apices. M. heterophyllum appeared to serve as a neutral substrate in terms of community composition but had a positive effect on the total numbers of algae.     

The response of stream periphyton to Pacific salmon: using a model to understand the role of environmental context

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Coal mine drainage effects on a lotic ecosystem in Northwest Colorado,U.S.A.

An aquatic biological survey was conducted in 1979–1980 to determine the effects of drainage from an active coal strip-mine on Trout Creek, in northwest Colorado, U.S.A. Sampling was conducted over four seasons at four stations for periphyton, benthic invertebrates and fish. Periphyton in Trout Creek changed in the relative abundance of algae divisions in no apparent relation to mining. Diatoms were the predominant division at all sites. Golden-brown algae were abundant in spring at the stations upstream and adjacent to the mine. Blue-green algae were relatively important at stations upstream and downstream of the mine in winter. Benthic invertebrates exhibited a progressive increase in density, biomass and number of taxa from the upstream station to the downstream station. Shannon-Wiener diversity index for bethic invertebrates decreased slightly downstream of mine drainage but remained indicative of a clean water community. Aquatic insects (especially Trichoptera) were the predominant invertebrates at all stations. Analysis of functional groups of benthic invertebrates revealed increased importance of collector species at the lower sites while shredders were most important upstream of the mine. Unlike the invertebrates, fish exhibited slightly lower biomass at the station adjacent to the mine. The decrease was due to fewer salmonids. However, salmonid density and biomass increased substantially at the station just downstream of the mine. Non-game species (suckers and minnows) increased in numbers downstream and were most abundant at the lowest station. This coal strip-mine had little discernable adverse effects on the periphyton and invertebrates of Trout Creek. Fish populations did not appear to be significantly affected by the mine. Apparently, the presence of settling ponds and a buffer zone of unmined land between the mine and the stream helped to minimize adverse effects.     

Interactions between neighboring algae and snail grazing in structuring microdistribution patterns of periphyton

The micro-distribution of periphyton (filamentous algae) on homogeneous substrates was examined in experimental tanks with and without the pressure of grazing snails. The growth of periphyton attached to artificial substrate was estimated at a small spatial scale (9.3 mm×9.3 mm cells) by varying the number of grazers (0, 5, or 10 snails per tank), using image processing analysis without removing the periphyton. The results suggest that periphyton growth within a cell was negatively affected by the biomass of periphyton in the cell but was positively affected by the biomass of periphyton in neighboring cells. A semivariogram analysis indicated that spatial heterogeneity increased with increasing grazing pressure. The size of patches was not clearly related to the number of snails, but there was a tendency for relative patch size to increase with snail density. Computer simulations were also conducted to examine factors affecting the degree of spatial heterogeneity. The simulation studies indicated that snails should graze a site that was previously grazed in order to produce the observed spatial heterogeneity of periphyton. The results also indicated that the positive effects of neighboring periphyton on the growth of algae might create patches. The interactions among neighboring algae and snail grazing might be an important factor creating the spatial heterogeneity of periphyton even on homogeneous substrates.     

The influence of flow and other environmental factors on benthic invertebrates in the Sacramento River, U.S.A.

We examined how community composition of benthic invertebrates was related to current velocities and other environmental variables within the Sacramento River in California, USA. Invertebrates were collected in 1998 and 1999 from 10 sites over a gradient of 187 river kilometers. Canonical correspondence analysis revealed that current velocity was the most important variable explaining community composition. Other predicator variables that influenced community composition included periphyton biomass, altitude, and disturbance. Because of the importance of velocity in structuring benthic communities in this system, alterations of flow caused by changes in river regulation structures should be carefully considered.     

Periphyton communities in a pristine mountain stream above and below heavy metal mining operations

SUMMARY. 1. Changes in species composition of the periphyton on introduced substrates were determined in an oligotrophic mountain stream subject to long-term heavy metal contamination.
2. At the upstream control site, the numerically most abundant taxa were Bacillarioph yta i( Achnanlhes minutissima, Achnanthes microcephala and Achnanthes linearis ) as well as, in summer, the Chlorophyta ( Mougeotia spp. and Ulothrix subtilissima ).
3. At the downstream contaminated site the periphyton community was totally dominated by Bacillariophyta throughout the sampling period. A, minutissima and A, microcephala were co-dominants during spring. Seasonal succession patterns did not parallel those at the upstream site. Chlorophyta were virtually absent and A. minutissima comprised 94% of the community during summer.
4. Species diversity, species evenness and dissimilarity index were utilized to detect differences in species composition, abundance and number. Slight differences were found in spring samples while summer samples indicated major differences between sampling sites.     

An ecologically useful classification of mean and near-bed flows in streams and rivers

SUMMARY. 1. Mean motion and near-bed flows in streams and rivers can be described using a classification derived from fairly simple field measurements. Our proposed classification is ecologically useful because it incorporates the combined effects of velocity, depth and substrate roughness to provide a means of quantifying the flow regimes occurring within the microhabitats of stream benthos. 2. Mean motion is characterized by the Reynolds number and the Froude number. Both are easily calculated, and because they are dimensionless they provide a means of comparing flows at different sites. 3. Five categories of near-bed flows (i.e. the flow microenvironments of stream benthos) are recognized. Flow may be hydraulically smooth or hydraulically rough and the latter category is subdivided further into: chaotic flow, wake interference flow, isolated roughness flow and skimming flow. Hydraulically smooth flows occur in sections of a river bed with fine sediments (e.g. sands, muds and clays). over flat sheets of bedrock, or in association with the flat blades of submerged macrophytes. Hydraulically rough flows occur where the substrate elements are larger (e. g. pebbles, cobbles and boulders) and are a function of substrate roughness and the depth of flow relative to the height of the roughness elements. Chaotic flows and wake interference flows predominate in riffles whilst isolated roughness flows and skimming flows are more likely to be a feature of runs. 4. Conventional stream sampling methods (e.g. the Surber and box or cylinder samplers) may collect across several different flow microhabitats. Our classification should enable different flow microenvironments to be recognized and so sampled more appropriately which, in turn, may reduce apparent clumping and the wide confidence intervals of benthic population estimates. Because our classification identifies ‘patches’ within the flow regime associated with the stream bed it enables stream ecologists to generate testable hypotheses regarding the distribution and abundance of benthic species in response to flow. 5. Our classification identifies spatial patterns in the flow regimes associated with the stream bed. Temporal patterns have not been identified: however, predictable changes in spatial patterns will resuh from temporal changes in stream discharge.     

Larval settlement in benthic environments: the effects of velocity and bed element geometry

1. Fluid‐mediated transport can play a key role in determining patterns of distribution and abundance for many benthic invertebrates. One critical challenge in understanding this process is to determine how flow patterns affect larval settlement, especially in those benthic environments where near‐bed flows interact with irregular bed topography to create complex variations in habitat suitability and settlement probability. 2. Boundary‐layer separation over topographical projections on an irregular bed can create two distinct regions of near‐bed flow (i.e., accelerating flow over the forebody and a zone dominated by slower eddies over the aftbody) that may have different effects on larval settlement. 3. We manipulated the flow over a convex roughness element (i.e., hemicylinder) in a flume and examined how the settlement of larvae of the black fly Simulium tribulatum varied with changes in near‐bed velocity and location over the substrate. Larval settlement rate was standardised to correct for variations in larval supply (i.e., among‐trial differences in the concentration of larvae in suspension). 4. Our analyses showed that position on the hemicylinder and near‐bed velocity both affected settlement rate, with a strong interaction effect. In particular, the observed relationship between settlement rate and velocity was negative on the substrate’s forebody and positive on the aftbody. We explore these results by considering potential physical and behavioural mechanisms affecting larval settlement. 5. The presence of a positive relationship between flow and settlement rate in the aftbody may allow settlement on bed elements in habitat where preferred fast‐flow conditions are present, but where settlement would otherwise by hydrodynamically limited. Thus, greater attention to settlement mechanisms in more realistic, topographically complex environments can not only help explain distribution patterns within substrates, but also among substrates and across habitats.