The influence of microhabitat on availability of drifting invertebrate prey to a net-spinning caddisfly

SUMMARY. 1. Invertebrate drift was sampled at both a rockface and a deep pebble riffle site in streams draining both a clearcut and a forested catchment.
2. A sampler was designed to separate the bottom 2 cm of flow, encompassing the effective range of caddisfly (Hydropsychidae: Trichoptera) catchnets. from upper flow.
3. No significant difference in drift density (numbers per cubie metre) was seen between sites within each stream. However, numbers per square centimetre intake area per day at the rockface sites were 4 times higher in the clearcut and 10 times higher in the forested stream than at the pebble-riffle site.
4. Rockface habitat which had highest drift availability was also the site of maximum secondary production of the predaceous collector- filterer Parapsyche cardis in both streams studied.
5. Increased sediment load in the clearcut stream may influence the efficiency of utilization of invertebrate drift by collector-filterers.     

Stream hydraulics as a major determinant of benthic invertebrate zonation patterns

SUMMARY. 1. Studies on the zonation of benthic fauna in fourteen streams situated in a variety of latitudes from Alaska to New Zealand have been evaluated.
2. We suggest that physical characteristics of flow ('stream hydraulics') are the most important environmental factor governing the zonation of stream benthos on a world-wide scale.
3. From the source to the mouth of a stream, zones of transition in stream hydraulics' occur, to which the general pattern of stream invertebrate assemblages can be related. In these zones benthic community stability and resilience must be different from those upstream and downstream of the hydraulic transition zones.     

Effects of habitat structure on habitat use by Gammarus pulex in artificial streams

1. The influence of coarse substratum and flow, coarse substratum and food, and predation risk and flow on habitat use by Gammarus pulex was studied in three experiments conducted in artificial stream channels. Each stream channel consisted of a riffle and pool habitat.
2. Location of coarse substrata and food was manipulated by placing cobbles (coarse substratum) and leaf packs (food) in different habitats. Predation risk was varied by running experiments in the presence and absence of sculpins ( Cottus gobio ), and flow was varied by pumping water with one or two pumps.
3. In all experiments Gammarus were most abundant in pools but placement of cobbles in riffles increased use of the latter. An even greater percentage of Gammarus used riffles if leaf packs were also placed there. Decreased discharge and the presence of sculpins ( Cottus gobio ) also caused Gammarus to increase use of riffles. These data indicate that Gammarus is able to evaluate differences in habitat quality and respond accordingly.     

Effects of habitat structure on habitat use by Gammarus pulex in artificial streams

1. The influence of coarse substratum and flow, coarse substratum and food, and predation risk and flow on habitat use by Gammarus pulex was studied in three experiments conducted in artificial stream channels. Each stream channel consisted of a riffle and pool habitat.
2. Location of coarse substrata and food was manipulated by placing cobbles (coarse substratum) and leaf packs (food) in different habitats. Predation risk was varied by running experiments in the presence and absence of sculpins ( Cottus gobio ), and flow was varied by pumping water with one or two pumps.
3. In all experiments Gammarus were most abundant in pools but placement of cobbles in riffles increased use of the latter. An even greater percentage of Gammarus used riffles if leaf packs were also placed there. Decreased discharge and the presence of sculpins ( Cottus gobio ) also caused Gammarus to increase use of riffles. These data indicate that Gammarus is able to evaluate differences in habitat quality and respond accordingly.     

Stream inflow and predation risk affect littoral habitat selection by benthic fish

1. We examined small, fishless headwater streams to determine whether transport of macroinvertebrates into the littoral zone of an oligotrophic lake augmented food availability for Cottus asper, an abundant predatory fish in our study system. We sampled fish and macroinvertebrates during the recruitment and growth season of 2 years, either monthly (2004) or bi‐monthly (2005), to observe whether stream inputs increased prey availability and whether variation in total macroinvertebrate biomass was tracked by fish. 2. Observations from eight headwater streams indicated that streams did not increase the total macroinvertebrate biomass in the shallow littoral zone at stream inflows, relative to adjacent plots without stream inputs (controls). The taxonomic composition of stream macroinvertebrates drifting toward the lake differed from that in the littoral lake benthos itself, although there was no evidence of any species change in the composition of the littoral benthos brought about by stream inputs. 3. Although streams made no measurable contribution to the biomass or taxonomic composition of the littoral macroinvertebrate benthos, there was substantial temporal variation in biomass among the eight sites for each of the (n = 7) sample periods during which observations were made. Variation in total biomass was primarily a function of bottom slope and benthic substrata in the lake habitats. Dominant taxonomic groups were Baetidae, Ephemerellidae (two genera), Leptophlebiidae, Chironomidae (three subfamilies) and Perlodidae, although we did not determine the specific substratum affinities of each taxon. 4. Mixed effects linear models identified a significant interaction between macroinvertebrate biomass and plot type (stream inflow vs. control) associated with fish abundance. Across the observed range of macroinvertebrate biomass, fish showed a significant preference for stream inflows, but more closely tracked food availability in the controls. For young‐of‐the‐year (YOY), a negative effect of temperature was also included in the model, and we observed lower temperatures at stream inflows. However, abundance of predatory adults affected habitat selection for YOY. Lake‐bottom slope also accounted for variation in abundance in both fish models. 5. Our results suggest that the effect of fishless headwater streams on downstream fish may not always be through direct delivery of food. In this study system, fish preferred stream inflow plots, but this preference interacted with macroinvertebrate biomass in a manner that was difficult to explain. For YOY, predation risk was related to the preference for stream inflows, although the specific factor that mitigates predation risk remains poorly understood.     

Associations between stream fish and benthos across environmental gradients in southern Ontario, Canada

1. The relationship between fish and benthic invertebrate communities in wadeable streams in southern Ontario, Canada, was examined using three independent and spatially distinct data sets. 2. Associations between fish and benthos were always significant when benthos were identified to family and often at the level of phylum. Identification to genus improved the strength of the fish–benthos association in one case. In contrast, identification to species did not improve the strength of the fish–benthos association. Associations between benthos and fish were weaker for one study utilizing a ‘rapid’ bio-assessment protocol involving field sorting and identification, and a second study which utilized winter benthic collections. 3. In two surveys, stream temperatures were important to the distributions of both fish and benthos, while in a third survey, fish and benthos were primarily influenced by stream size. In all three studies, fish and benthos were associated with similar suites of environmental variables, suggesting that the fish–benthos associations in these streams was driven by corresponding environmental tolerances. 4. Although there was significant variation in the strength of the fish–benthos association which could be attributed to differences in sampling methodologies, the findings from the present study confirm that stream fish and benthos are significantly associated. Therefore, surveys of benthos can be used to make inferences on the condition of fish community composition.     

Indicating shear stress with FST-hemispheres - effects of stream-bottom topography and water depth

1. In order to quantify the effects of substratum roughness on shear stress and Fliesswasserstammtisch-(FST)-hemisphere movement, hemispheres were calibrated against shear stress in a laboratory flume with fully developed turbulent flow. In five different runs, substratum roughness, water depth and location of hemispheres in relation to the surrounding particles, were varied. 2. FST-hemisphere results were strongly influenced by bottom topography. In the case of hydraulically rough flow a linear relationship exists between shear stress (χ_c) and hemisphere density (ρ_h), whereas in the case of quasi-smooth flow a power law was obtained for the χ_c/ρ_h, relationship. Shear stress for a given hemisphere and relative roughness h/k > 4 (h= water depth; k= height of the roughness elements) deviated up to one order of magnitude between roughnesses. In water depths, where h/k 4, the χ_c/ρ_h relationships are dependent on the ratio h/k, due to water surface effects on hemisphere movement. In the case of k = d₉₀=r_h (d₉₀= characteristic diameter of the largest particles of the bottom substratum; r_h= radius of the hemispheres), the location of the hemispheres in respect to the roughness elements is of secondary importance. 3. In the case of hydraulically rough flow and wake interference between the roughness elements, the turbulent flow field close to the substratum is three-layered, each layer being characterized by its own velocity distribution laws (Dittrich & Hammann de Salazar, 1993). Depending on the height of the roughness elements, FST-hemispheres will be subjected mainly to flow forces of the near-bed layer (in rough substrata) or to flow forces distant from the near-bed zone (in fine substrata). The dominant flow forces acting on bottom particles, organisms, or FST-hemispheres are shear force and lift force. 4. The Local Shear Stress Model (Lamouroux et al., 1992) leads to a correct prediction of hemisphere distribution in a stream with a cobble size substratum, but to an overestimation of hemisphere numbers in a sandy-bottom stream. The substratum-dependent shear stress values therefore need to be entered into the model and a measure of substratum roughness included. 5. Macroinvertebrate abundance correlates well with the movement of FST-hemispheres. Samples from points with very high or very low roughness did not contribute to scatter in the data, indicating that the sum of the near-bed flow forces is relevant to macroinvertebrate distribution, not shear stress alone. We conclude, that FST-hemispheres are well suited to characterize near-bottom hydraulics and therefore the microhabitat of the benthos.     

A continuous study of the total drift of freshwater shrimps, Gammarus pulex, in a small stony stream in the English Lake District

1. The objective was to determine the major factors affecting the downstream dispersal (drift) of freshwater shrimps, Gammarus pulex. Sample replication and frequency are major problems in the quantification of drift. For the first time, these problems were avoided by sampling the whole stream continuously so that all the shrimps drifting downstream at the sampling point were caught in a net emptied at dusk and dawn in 1966, and every 3 days in 1967. 2. There was no consistent seasonal pattern in drift rates, but a high proportion of annual drift was taken in only a few samples. There was a nocturnal diel pattern of drift with peaks soon after dusk and just before dawn. A power function described the significant (P < 0.001) relationship between drift and flow, and was used to neutralise the dominant effects of flow by standardising total drift over 24 h, nocturnal drift and diurnal drift (drift per 50 m³). These were all significantly (P < 0.001) related to benthos density, but not to date, temperature, or length of the night or day. 3. The relationship between drift values and the independent variables, flow and benthos density, was well described (P < 0.001) by a multiple‐regression model. Adding temperature, date, and/or the length of the night or day did not improve model fit. Variations in flow and benthos density explained 94% of the variation in total drift over 24 h, 97% of the variation in nocturnal drift, but only 44% of the variation in diurnal drift. A power function described (P < 0.001) the relationship between total drift and the volume of water sampled over 3‐day periods in 1967. Flow explained 95% of this drift variation; it was unnecessary to add another variable such as benthos density. 4. The significance of this study is that it avoided the problems associated with the quantification of drift samples. Therefore, the conclusions are more robust than those of many previous studies. A high proportion of the annual drift losses would have been undetected by intermittent sampling. Temperature, season, night or day length had no significant effect on drift densities, and the relationship between drift and benthos densities was proportional, not density dependent. The nocturnal increase in drift could not be interpreted as an antipredator behaviour. The dominance of flow and benthos density was apparent but the quantitative relationships posed further questions, especially those related to drift distances at different velocities.     

A comparison of bacteria and benthic invertebrates as indicators of ecological health in streams

1. We set out to evaluate the reliability of bacterial communities as an indicator of freshwater ecological health.
2. Samples of epilithic biofilm were taken over a 1-year period from four streams, each impacted by varying degrees of human modification. The bacteria within each sample were characterised using a whole community DNA fingerprinting technique (automated ribosomal intergenic spacer analysis). Spatial and temporal differences in community structure between samples were visualised using multi-dimensional scaling and quantified using permutational multivariate anova . Macrobenthic invertebrates, which are commonly used as indicators of stream ecological health, were also sampled for comparison.
3. Multivariate analysis revealed a clear gradient in macroinvertebrate community structure between sites exposed to increased human impact. Bacterial communities, however, could only distinguish the most impacted site from the remainder.
4. Additional research is required to increase the sensitivity of bacterial community analyses before endorsing their use as an indicator of freshwater ecological health.     

Macrozoobenthos communities and biocoenotic assessment of ecological status in relation to degree of human impact in small streams in southwest Germany

One of the major objectives of the VALIMAR project is to determine the ecological significance of various fish biomarker studies as indicators of chronic pollution in small streams in southwest Germany. Results of these fish biomarker investigations were compared to information from complementary studies on the meiobenthos, macrobenthos, and fish community studies to assess the ecological significance of these biomarker investigations. The main objective of this study was to provide biological assessments of the biomarker sites on the basis of the macrozoobenthos communities. Since no validated framework for the assessment of the biological integrity existed in Germany, two multimetric approaches were adapted to the whole stream system by investigating benthos and fish communities of 46 sites of varying degrees of human disturbance. Assessment of the communities was conducted in accordance with the European Community Water Framework Directive. Species distribution of benthos depended upon stream type and pollution status of streams. Biological attributes and bioindices of benthos communities, however, did not correlate with typological parameters like stream size or dominant substrate but correlated better with pollution parameters like conductivity or chloride concentration. Using a set of 18 measures, such as portion of sessile individuals, Rheoindex, oxygen availability index, and portion of pool dwellers, the benthos communities were characterized and evaluated. The composition of the fish communities was mainly determined by stream type, pollution and migration barriers. The influence of chemical parameters could be assessed by developing a“fish chemistry index”, which calculatesthe similarity of the present fish community with the potential natural community, but excludes those species strongly effected by deficits in stream channel morphology. Both fish chemistry index and benthic indices strongly correlated with pollution index parameters, clearly distinguishing between the more polluted Körsch sites and the less pollutedKrähenbach and Aich sites. Most of the single bioindices as well as overall assessment by multimetric indices indicated a gradient of decreasing quality from the reference stream Krähenbach to theslightly polluted Aich and Körsch upstream site (KE, upstream of all sewage treatment plants) and finally to the most polluted Körsch site directlybelow the most upstream sewage treatment plant (KD). According to the Water Framework Directive, the classification of ecological status of the benthos communities ranges from “high” (best of 5 classes) forthe reference stream to “bad” (5th class) for KD. Assessment of the fish community tends to score somewhat worse than the benthos due to deficits in morphological quality of the stream reaches. The benthos assessment and the newlydeveloped “fish chemistry index” correlated well with chemical water quality and hence with biomarkers, whereas ecological status of fish and overall ecological status was also influenced by river morphology. In conclusion all tested assessment methods on biocoenotic level are reliable indicators for the degree of human disturbance on small streams, whereas biomarkers are more suited for risk assessment and the investigation ofcause-effect-relationships.     

Freshwater ecosystem structure–function relationships: from theory to application

1. In this issue we aimed to answer the questions: (i) under what circumstances are functional variables better than structural ones for assessing ecosystem health? and (ii) are there good indicators of change in ecological functioning along perturbation gradients?
2. Of the numerous functional indicators tested in this issue, several show a response to anthropogenic stress and could be included in assessments of ecosystem health and integrity in running waters.
3. In three of eight studies, function showed a stronger response to anthropogenic stress than structure, whereas one study showed a response in structure and not function, and four studies showed responses in both structure and function. Thus structure alone could not detect all types of impairment and functional aspects should also be included and further developed for assessing running-water ecosystem health and integrity. Functional variables may be especially useful in situations where there is a stronger response among organisms not usually included in stream assessment (e.g. fungi and bacteria) than the commonly used invertebrate, macrophytes and fish indicators.
4. Leading research questions related to the use of functional indicators in running waters include: (i) how large is natural and operator-induced variation for functional indicators? (ii) how small of an effect size (delta) can be detected using structural versus functional indicators? and (iii) how do we efficiently improve theories as well as predictive ability for functional measures to assess the effects of anthropogenic stressors?
5. To advance the use of functional indicators in applied running-water studies, we need to supplement the approach of using large-scale datasets and correlation with ecosystem manipulations.     

Calibration of FST-hemispheres against bottom shear stress in a laboratory flume*

SUMMARY.

• 1 Fliesswasserstammtisch (FST)-hemispheres of identical size but different densities were exposed on a horizontal plane on the pebble-covered bottom of a laboratory flume at streaming to shooting, turbulent and fully developed flow. The heaviest hemisphere moved was used as an indicator of shear stress, a flow force acting on the flume bottom.
• 2 The relationship between shear stress and hemisphere density was practically the same for three different sets of hemispheres, namely one prototype set newly prepared for the calibrations, one prototype set intensively used in the field prior to calibrations, and one randomly selected new set bought commercially.
• 3 Shear stress (y, dyn cm^?2) was related to hemisphere density (x, g cm^?3) by y= x^2.85 (n= 69; r²= 0.993). For lighter hemispheres this relationship was better explained by the linear model y= 7.32x– 6.60 (n= 35; r²=0.983).

     

The use of photosynthesis inhibitor (DCMU) for in situ metabolic and primary production studies on soft bottom benthos

A selective chemical photosynthesis inhibitor, DCMU (Dichorophenyl-dimethylurea), dissolved in DMSO (Dimethyl sulfoxide) was substituted for the dark incubation method commonly used to measure the oxygen consumption in metabolic and primary production studies. We compared oxygen fluxes during light incubations with DCMU and dark incubations procedure, on soft bottom benthos. For this purpose, we studied the effects of different DCMU concentrations. A concentration of 5 · 10^–5 mol l^–1 inside a clear incubation enclosure completely inhibits photosynthesis without affecting the metabolism of soft bottom benthos.     

Blood flow in the brain and liver of chicken in embryonal and early postembryonal periods

In chicken Leghorn, blood flow volume speed (BF, laser-Doppler flowmetry) in the brain hemispheres and in liver was measured on days 10, 14, and 19 of embryogenesis and on day 4 after hatching (in experiments on late embryos and chickens, urethane narcosis was used). It was revealed, that initial BF in investigated organs was 2-fold lower than earlier measured in skeletal muscles. In the liver, low BF remained at all periods, but it grew 5-fold greater after hatching. In the brain hemispheres, the BF during this period grows gradually reaching 4-fold size in chickens. It was shown that blood stream increase in the brain was accompanied by uniform increase in anatomic lumen of internal carotid artery; thus settlement sizes of linear speed of blood flow and wall shear stress remain in it at the same level. Lumen extension of celiac artery during the observation period lags behind increases in a blood stream of in it that leads to increase in it of the named parameters.     

Field experiments on the mobility of benthic invertebrates in a southern English stream

1. In each of twenty-six week-long experiments, the colonization by macroinvertebrates of boxes of natural sediment in a stony stream was measured. The experiments took place between February and November 1992 and environmental conditions prevailing during the weeks (particularly discharge and temperature) differed widely.
2. Colonization rate also varied widely between the weekly experiments and was sensitive to discharge, temperature and background benthic density, depending on the taxa considered.
3. A 'mobility index' measured colonization rate independently of background benthic density. This index was most strongly and positively correlated with discharge for the abundant stonefly Leuctra nigra, the net-spinning caddis Plectrocnemia conspersa and the Chironomidae, perhaps indicating that their mobility is physically driven by the influence of flow on drift.
4. Mobility in a second stonefly ( Nemurella pictetii ) was greatest in summer, when flows were low but temperature was high. Peak mobility in this species occurred during an ochreous bloom on the stream bed. It is likely the mobility of N. pictetii is more active than the other taxa.
5. There were thus differences among taxa in their mobility at baseflow and in their susceptibility to flow fluctuations. The population consequences of differences in mobility among taxa are discussed.     

The effect of the acanthocephalan parasite (Pomphorhynchus laevis) on the drift of its intermediate host (Gammarus pulex)

SUMMARY 1. The drift of Gammarus pulex in a population in which approximately 20% of adults are infected with the acanthocephalan parasite Pomphorhynchus laevis was monitored at margin and mid-river sites in the River Teme, England over a 24-h period in mid-summer.
2. Drift densities showed a diurnal pattern with a large increase at night, independent of parasite burden.
3. There was no significant effect of site (margin or mid-river) on the proportion of parasitized and unparasitized G. pulex found in the drift or the benthos.
4. The drift of parasitized G. pulex was significantly greater than unparasitized animals. Gammarids harbouring only one parasite were found in significantly higher proportions in the drift than those with two or more parasites.
5. At both sites (margin and mid-river) the proportion of unparasitized adult G. pulex in the drift was significantly lower than that in the benthos. However, there was a significantly higher proportion of parasitized animals in the drift than in the benthos.     

Influence of ultraviolet-B radiation on the drift response of stream invertebrates

1. Recent studies have shown that ultraviolet (UV) radiation (280–400 nm) has increased by ≈ 8% in temperate regions over the past decade, but little effort has been devoted to understanding the ecological effects on temperate ecosystems. This research examined the effects of artificial ultraviolet-B (UVB; 280–320 nm) radiation on the drift response of immature stream insects in laboratory microcosms.
2. Two experiments involved natural populations of stream invertebrates, collected from the Cache la Poudre River (September 1994) and the Arkansas River (October 1995) in Colorado. UVB lamps were turned on from 10.00 to 14.00 h each day, and drifting animals were collected on days 1, 3, 5 and 7 during the exposure period. Levels of artificial UVB used in these experiments were similar to levels that stream organisms experience during clear, mid-day conditions at Fort Collins, Colorado (longitude 105°30'; latitude 40°35').
3. Drift was significantly higher in microcosms exposed to UVB than in controls and was dominated by Baetis sp. (Ephemeroptera: Baetidae), Trichoptera (caddisflies) and Simulium sp. (Diptera: Simuliidae). The increased drift of some stream invertebrates in UVB-exposed streams may be a behavioural response and/or a result of injury.
4. Stream organisms may be particularly sensitive to predicted increases in UV radiation, because streams are generally shallow with clear water. As a result of this potential sensitivity, we recommend that research be directed to understanding the ecological effects of UV radiation on these habitats.     

Influence of ultraviolet-B radiation on the drift response of stream invertebrates

1. Recent studies have shown that ultraviolet (UV) radiation (280–400 nm) has increased by ≈ 8% in temperate regions over the past decade, but little effort has been devoted to understanding the ecological effects on temperate ecosystems. This research examined the effects of artificial ultraviolet-B (UVB; 280–320 nm) radiation on the drift response of immature stream insects in laboratory microcosms.
2. Two experiments involved natural populations of stream invertebrates, collected from the Cache la Poudre River (September 1994) and the Arkansas River (October 1995) in Colorado. UVB lamps were turned on from 10.00 to 14.00 h each day, and drifting animals were collected on days 1, 3, 5 and 7 during the exposure period. Levels of artificial UVB used in these experiments were similar to levels that stream organisms experience during clear, mid-day conditions at Fort Collins, Colorado (longitude 105°30'; latitude 40°35').
3. Drift was significantly higher in microcosms exposed to UVB than in controls and was dominated by Baetis sp. (Ephemeroptera: Baetidae), Trichoptera (caddisflies) and Simulium sp. (Diptera: Simuliidae). The increased drift of some stream invertebrates in UVB-exposed streams may be a behavioural response and/or a result of injury.
4. Stream organisms may be particularly sensitive to predicted increases in UV radiation, because streams are generally shallow with clear water. As a result of this potential sensitivity, we recommend that research be directed to understanding the ecological effects of UV radiation on these habitats.     

Field experiments on flow refugia in streams

1. Field experiments were carried out to determine whether animals move into areas of low shear stress during periods of peak flow and therefore avoid hydraulic disturbance.
2. Flow at the scale of 0.05 m² patches was reduced experimentally by creating artificial 'refugia'. Invertebrate colonization of cages with 1.1 mm mesh sides, which provided such potential refugia by reducing hydraulic forces within the cage, was compared with colonization of cages with coarser (15 mm) mesh which did not restrict the flow.
3. Colonization of these cages was tested over a series of weekly periods in two different streams. Nine trials were completed in a stream with abundant natural flow refugia (Broadstone Stream, SE England), and during three of these trials strong spates occurred. Six trials were completed in a stream with comparatively few natural flow refugia (Dargall Lane, SW Scotland), and peak flows were relatively less.
4. In Broadstone Stream, the relative colonization of refugium cages was greatest during periods of peak flow, suggesting animals had used these low-flow areas as refugia during spates. Use of the artificial refugia did not occur in Dargall Lane, at least at the flows achieved during the trials.
5. Our results are consistent with the hypothesis that stream invertebrates accumulate in refugia during high flow disturbances. Whether their distribution among patches is altered by active or passive means remains unclear.     

Zebra mussels mediate benthic–pelagic coupling by biodeposition and changing detrital stoichiometry

1. The zebra mussel ( Dreissena polymorpha ) is one of the most successful invasive species; it has colonised many aquatic systems in Europe and North America with strong impacts on various ecosystem processes. The effect of D. polymorpha filtration on pelagic seston concentrations has been quantified in several studies, but the magnitude and stoichiometry of the transfer of sestonic biomass into benthic detritus by D. polymorpha and the accompanying enrichment of the benthic habitat is still under-investigated.
2. We studied biodeposition by zebra mussels in two series of laboratory experiments with the food algae Cryptomonas erosa and Scenedesmus obliquus . We also measured the year-round biodeposition rate under natural conditions in the oligotrophic Lake Constance.
3. In all experiments, zebra mussel biodeposition was linearly related to seston concentration. In the field, the relationship changed with a seasonal shift in algal composition and lower biodeposition rates during the spring algal bloom.
4. For both algal species in laboratory experiments, biodeposited material was depleted in phosphorous at an algal concentration ≤0.6 mg ash-free dry mass L⁻¹, but not at higher concentrations. This effect was not observed in the field, probably because of high variation in C : N : P stoichiometry.
5. By mediating the transfer of pelagic resources into the benthos zebra mussels provide a sufficient amount of detritus for benthic invertebrates, especially during summer. Thus, material biodeposited by the mussels might increase benthic secondary production from pelagic resources, and zebra mussels are important mediators of this flux of organic matter from the pelagic zone into the benthos.