The relative importance of drift causes for stream insect herbivores across a canopy gradient

A key attribute of riverine food webs is the downstream movement of invertebrates via the water column, or invertebrate drift. Causes of drift include benthic predation, food limitation, and perhaps passive entry, which may occur when invertebrates lose their purchase on stream substrate. However, the relative importance of drift causes is unknown, as is whether the relative importance of drift causes varies across space. Combining observational data on invertebrate herbivore and predator guild densities with in‐stream experiments, we evaluated the relative importance of benthic predation, food limitation, and passive entry as proximate causes of drift for the herbivore guild across the canopy gradient of a montane stream. We found that 1) benthic predation and food limitation were both more important as causes of herbivore drift than passive entry; 2) drift caused by food limitation did not vary with riparian canopy, whereas herbivore density decreased with increasing riparian canopy, and 3) per capita drift increased linearly with increasing density, while per capita drift decreased in a negative hyperbolic fashion with increasing food, indicating that herbivore drift is proportional to herbivore density, and inversely proportional to food. We conclude that invertebrate herbivore drift was overwhelmingly an active process to improve fitness, and that herbivore food did not vary across the canopy gradient, likely because increased herbivory from larger herbivore populations at sunnier sites prevented food from accumulating.     

Predation and drift of lotic macroinvertebrates during colonization

Summary A field experiment was carried out to determine the effect of an invertebrate predator on the colonization and drift of benthic macroinvertebrates in experimental stream channels. Lotic invertebrates colonized four replicate channels: two controls with no predators, and two channels with low densities (2.8 m^–2) of predatory stonefly nymphs, Doroneuria baumanni (Perlidae). Immigration rates were measured at the inflow of two other channels. Drift rates of invertebrates immigrating to and emigrating from channels were measured daily, and benthic samples were collected every five days. Over a 25-day colonization period, benthic densities of Baetis nymphs and larval Chironomidae were reduced by D. baumanni. Colonization curves were fit with a power function and significantly different colonization rates were indicated for both Baetis and chironomids in predation and control channels. A predator-induced drift response was exhibited by Baetis only and this response was size-dependent. In the presence of D. baumanni, large Baetis drifted more frequently than small nymphs and, correspondingly, small nymphs were more frequent in the benthos. Net predator impacts on invertebrate densities in channel substrates were partitioned into predator-induced drift and prey consumption. These estimates suggest that predator avoidance by Baetis is a prominent mechanism causing density reductions in the presence of predators. Reductions in the density of Chironomidae, however, were attributed to prey consumption only. A rainstorm during the experiment demonstrated that stream flow disruptions can override the influence of predators on benthic invertebrates, at least temporarily, and re-set benthic densities.     

Habitat effects on invertebrate drift in a small trout stream: implications for prey availability to drift-feeding fish

In this study, we focused on the drivers of micro- and mesohabitat variation of drift in a small trout stream with the goal of understanding the factors that influence the abundance of prey for drift-feeding fish. We hypothesized that there would be a positive relationship between velocity and drift abundance (biomass concentration, mg/m³) across multiple spatial scales, and compared seasonal variation in abundance of drifting terrestrial and aquatic invertebrates in habitats that represent the fundamental constituents of stream channels (pools, glides, runs, and riffles). We also examined how drift abundance varied spatially within the water column. We found no relationship between drift concentration and velocity at the microhabitat scale within individual pools or riffles, suggesting that turbulence and short distances between high- and low-velocity microhabitats minimize changes in drift concentration through settlement in slower velocity microhabitats. There were also minimal differences in summer low-flow drift abundance at the mesohabitat scale, although drift concentration was highest in riffle habitats. Similarly, there was no differentiation of drifting invertebrate community structure among summer samples collected from pools, glides, runs, and riffles. Drift concentration was significantly higher in winter than in summer, and variation in drift within individual mesohabitat types (e.g., pools or riffles) was lower during winter high flows. As expected, summer surface samples also had a significantly higher proportion of terrestrial invertebrates and higher overall biomass than samples collected from within the water column. Our results suggest that turbulence and the short length of different habitat types in small streams tend to homogenize drift concentration, and that spatial variation in drift concentrations may be affected as much by fish predation as by entrainment rates from the benthos. Handling editor: Robert Bailey     

Intra- and interspecific dominance hierarchies and variation in foraging tactics of two species of stream-dwelling chars

Aggressive interactions, foraging behavior, habitat use and diet were studied in sympatric populations of white-sported char,Salvelinus leucomaenis, and Dolly Varden,Salvelinus malma, in a Japanese mountain stream. Underwater observations on individuals of both species revealed two distinct behavioral regimes: aggressive drift foragers and non-aggressive benthos foragers. Aggressive drift foragers defended partial territories around focal points from which they made forays to capture invertebrates drifting in the water column. Non-aggressive benthos foragers cruised around and beneath cobble in large foraging ranges that overlapped each other. Intra- and interspecific, size-dependent dominance hierarchies were recognized among aggressive drift foragers, whereas non-aggressive benthos foragers showed no such relationships. Terrestrial invertebrates were the most abundant prey in the diets of drift foragers, whereas a very small proportion of the diet of benthos foragers was made up of these taxa. Benthos foragers showed more complex diet composition than drift foragers. These results suggest that non-aggressive benthos foragers may avoid not only interference but also exploitative competition by using alternative foraging tactics. The proportion of drift foragers to benthos foragers among white-spotted char was more than 35 times that among Dolly Varden. The significant difference in the proportion of each species using the two types of foraging strategy results in interspecific food segregation in sympatric populations.     

Crowded waters: short-term response of invertebrate drift to water abstraction

Water abstraction modifies the environmental conditions of stream ecosystems, which can affect invertebrate assemblages by altering drift. We examined this issue with a before–after–control–impact design experiment in which we diverted 90% of the natural flow from a headwater stream. We measured flow-reduction effects on drift densities (animals/m³), total drift rates (animals leaving the reach per hour) and net balance of invertebrates entering or leaving the Impact reach. We also identified the specific taxa and traits that drove these responses. The sudden decrease in flow promoted a 12-fold increase in overall drift density at the Impact reach, which was primarily driven by filterers, shredders and taxa associated with fast velocities, such as simulids. By contrast, drift densities of other abundant taxa, such as Baetis, Esolus and chironomids, increased less than what could be expected from the magnitude of flow reduction. While drift rates remained unchanged after water abstraction, the Impact reach became a net invertebrate exporter indicating that many taxa drifted actively as a response to stressful conditions rather than passively, which would be reduced by water abstraction. Therefore, our results suggest that water abstraction influences drift, with potentially important consequences for the invertebrate assemblages and ecosystem processes further downstream.     

Composition and seasonality of benthic invertebrates,and drift in the Naro Moru River,Kenya

A survey was carried out to establish the nature and composition of the benthos along the Naro Moru, a tropical river in central Kenya using artificial substrate baskets, from November 1986 to October 1987. A clear longitudinal zonation existed for Diptera and Ephemeroptera which were the major benthic taxa. Maximum colonization took place after ten days of exposure. Seasonal variations in abundance were also observed. All taxa collected from the bottom samples were also collected in the drift samples, but the percentage composition of the benthos showed variations with that of the drift. Simulium sp. dominated the benthos whilst Baetis spp. dominated the drift. There was a positive correlation between drift rate and benthic fauna density.     

Benthivorous fish reduce stream invertebrate drift in a large-scale field experiment

Drift as a low-energy cost means of migration may enable stream invertebrates to leave risky habitats or to escape after encountering a predator. While the control of the diurnal patterns of invertebrate drift activity by fish predators has received considerable interest, it remains unclear whether benthivorous fish reduce or increase drift activity. We performed a large-scale field experiment in a second-order stream to test if invertebrate drift was controlled by two benthivorous fish species (gudgeon Gobio gobio and stone loach Barbatula barbatula). An almost fishless reference reach was compared with a reach stocked with gudgeon and loach, and density and structure of the invertebrate communities in the benthos and in the drift were quantified in both reaches. The presence of gudgeon and stone loach reduced the nocturnal drift of larvae of the mayfly Baetis rhodani significantly, in contrast to the findings of most previous studies that fish predators induced higher night-time drift. Both drift density and relative drift activity of B. rhodani were lower at the fish reach during the study period that spanned 3 years. Total invertebrate drift was not reduced, by contrast, possibly due to differences in vulnerability to predation or mobility between the common invertebrate taxa. For instance, Chironomidae only showed a slight reduction in drift activity at the fish reach, and Oligochaeta showed no reduction at all. Although benthic community composition was similar at both reaches, drift composition differed significantly between reaches, implying that these differences were caused by behavioural changes of the invertebrates rather than by preferential fish consumption. The direction and intensity of changes in the drift activity of stream invertebrates in response to the presence of benthivorous fish may depend on the extent to which invertebrate taxa can control their drifting behaviour (i.e. active versus passive drift). We conclude that invertebrate drift is not always a mechanism of active escape from fish predators in natural streams, especially when benthos-feeding fish are present.     

Invertebrate responses to short-term water abstraction in small New Zealand streams

1. Small permanent streams are coming under increasing pressure for water abstraction. Although these abstractions might only be required on a short‐term basis (e.g. summer time irrigation), the highest demand for water often coincides with seasonal low flows. 2. We constructed weirs and diversions that reduced discharge in three small streams (<4 m width) to test the hypotheses that short‐term water abstractions would decrease habitat availability and suitability for invertebrates, resulting in increased invertebrate drift, reduced taxonomic richness and decreased benthic invertebrate densities. 3. We sampled benthic invertebrates, invertebrate drift and periphyton at control (upstream) and impact (downstream) sites on each stream before and during 1 month of discharge reduction. 4. Discharge decreased by an average of 89–98% at impact sites and wetted width decreased by 24–30%. Water depth decreased by 28–64% while velocity decreased by 50–62%. Water conductivity, temperature and dissolved oxygen showed varying responses to flow reduction among the three streams, whereas algal biomass and pH were unaffected in all streams. 5. The densities of invertebrate taxa tended to increase in the impact reaches of these streams, even though invertebrate drift increased at impact sites in the first few days following discharge reduction. There were a higher proportion of mayflies, stoneflies and caddisflies at the impact site on one stream after flow reduction. There were no changes to the number of taxa or species evenness at impact sites. 6. Our results suggest that for these small streams, the response of invertebrates to short‐term discharge reduction was to accumulate in the decreased available area, increasing local invertebrate density.     

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.     

Use of both benthic and drift sampling techniques to assess tropical stream invertebrate communities along an altitudinal gradient, Costa Rica

1. Two sampling techniques were used to characterize invertebrate communities in eight, low-order streams along an altitudinal gradient in Costa Rica that represents the last continuous tract of primary forest spanning such extremes in elevation (i.e. near sea level to 2900 m a.s.l.) along the Caribbean Slope of Central America. A standard Surber sampler was used to sample invertebrates on the stream bottom, and drift sampling nets were used to sample invertebrates drifting in the stream flow. 2. Sites were established at 30, 50, 700 1800 and 2700 m a.s.l. In one to two streams per site, six Surber samples were collected, and drift was sampled every 3 h over one 24-h period between April and August 1994. All sites were in primary forest, with the exception of the lowest elevation site (30 m) which was located in banana plantations. 3. Both sampling techniques indicated that Diptera (Chironomidae) and Ephemeroptera were the dominant insect groups at all sites. Disturbed streams draining banana plantations were dominated by Chironomidae and had lower taxon richness and diversity than other sites. 4. While data from benthic samples indicated that insects were the major faunal component (> 90%) at all sites, drift samples were dominated by larval shrimps (> 50%) at the 30 m and 50 m sites. 5. Drift periodicity of invertebrates was observed at those sites characterized by predaceous fishes: nocturnal drift densities were higher than diurnal densities at 30, 50 and 700 m a.s.l., however, no periodicity was observed at 1800 and 2700 m a.s.l. where fish were absent. 6. This study shows the importance of measuring invertebrate drift, in addition to directly sampling the benthos. Drift sampling provided data on a major community component (shrimps) of lowland tropical streams, that would have been overlooked using traditional benthic sampling techniques, and in some cases provided additional information on taxon richness. 7. Based on results of the present study, it is recommended that drift sampling be included as a standard complementary tool to benthic sampling in biological assessments (e.g. bioassessment protocols) of tropical streams, which are often characterized by migratory invertebrate species such as shrimps. Drift samples provide critical information on the presence or absence of shrimps and also on the timing and magnitude of their migration which is an important link between many tropical rivers and their estuaries.     

Ontogenetic changes in the drifting of four species of elmid beetles elucidate the complexity of drift-benthos relationships in a small stream in Northwest England

1. This study aimed to quantify ontogenetic changes in the drifting of Elmis aenea, Oulimnius tuberculatus, Esolus parallelepipedus and Limnius volkmari (Coleoptera: Elmidae), and to relate their drift to benthic density. Monthly samples were taken over 39 months, using three surface nets at each of two contrasting sites in a small stream: one in a deep section with abundant macrophytes, and the other in a shallow stony section. 2. Most larvae and adults were taken in the drift at night with little variation between catches in the three nets at each site. Day catches were very low, often zero. No significant relationships could be established between mean numbers in the drift catches and benthic densities. 3. When night catches were converted to drift densities (number caught per 100 m³ of water sampled), the latter were positively related to monthly losses in the benthos, but not to benthic densities. A linear regression described the relationship, and equations for the different life‐stages within each species were not significantly different from the equation for all life‐stages combined. However, drift losses were only about 0.07% of total losses in the benthos. A severe spate in October 1967 increased the number of larvae and adults in the drift, but not drift densities, except for immature adults of E. aenea, O. tuberculatus and E. parallelepipedus. 4. Key life‐stages with the highest drift density were the earliest life‐stage soon after egg hatching for E. aenea, the start of the larval overwintering period for O. tuberculatus and L. volkmari, and mature adults during the mating season for all three species. Drift density for E. parallelepipedus was too low to identify a key life‐stage. These key life‐stages corresponded with critical periods for survival in the life cycle, as identified in an earlier study in the same stream. Mortality was high during these critical periods, hence the strong relationship between drift density and benthic losses. The latter relationship was very consistent for different life‐stages within each species, and partially supported the rarely‐tested hypothesis that drift represents surplus production in the benthos.     

Effects of an experimental acid pulse on invertebrates in a high altitude Sierra Nevada stream

The effects of pulsed acidification on invertebrate densities and drift, and water chemistry, in a high altitude Sierra Nevada stream were measured using artificial stream channels. Water was diverted from the Marble Fork of the Kaweah River, California, U.S.A., through twelve replicate channels; however, low flow in the summer of 1985 eliminated all but four of these channels. Channels were stocked with natural substrates and organisms from the Marble Fork of the Kaweah River. After a three week acclimation period, we simulated a low pH rain event by adding acid (H₂SO₄ and HNO₃) to two of the channels, reducing pH to 5.0 for 6 hours. The other two channels acted as controls (pH 6.4). During acid additions, Baetis spp. drift in acidified channels was ca. 7 times higher than in control channels (F = 39.02, p < 0.025; data fourth root transformed, ANOVA), and the percentage of drifting baetids that was dead was significantly higher in acidified than control channels (46% vs. 0%, F = 29.86, p < 0.05; arcsine square root transformed data, ANOVA). Other taxa showed no significant drift responses, and benthic densities of all taxa showed no effects two days after acidification, probably owing to rapid recolonization by invertebrate drift in influent waters. Stream chemistry data are presented; heavy metal concentrations did not significantly increase in the 2 m stream channels.     

Invertebrate drift along a longitudinal gradient in a Neotropical stream in Serra do Cipó National Park,Brazil

The diversity and composition of drift invertebrate assemblages were evaluated along a longitudinal gradient of an altitudinal stream in southeastern Brazil. The main goal of this study was to evaluate the influence of seasonality, stream order, and some abiotic factors on invertebrate drift and the use of drifting invertebrate assemblages to assess aquatic invertebrate diversity. Drift samples were collected over a 24 h period using nets (open area of 0.08 m²; mesh 0.250 mm), partially submerged (60%) in the water column. Taxonomic richness, Pielou evenness (J), Shannon–Wiener diversity (H), and total density of drift invertebrate assemblages were used in unpaired t-tests, Kruskal–Wallis and stepwise multiple regression analysis. The results showed a high taxonomic richness of aquatic invertebrates, with 91 taxa found. Chironomidae and Ephemeroptera represented together c. 80% of the total density of drift organisms. The drift approach allowed the collection of new and rare taxa, besides the knowledge of pupae stage of several chironomid genera. Significant differences in the taxonomic richness and diversity of drift invertebrate assemblages were found between the rainy and dry periods, indicating a significant influence of seasonality. An increase in water flow and electrical conductivity were associated with the increase in the taxonomic richness and diversity in the rainy period. No significant differences were found among the other abiotic variables among the stream orders.     

Feeding modes in stream salmonid population models: is drift feeding the whole story?

Drift-feeding models are essential components of broader models that link stream habitat to salmonid populations and community dynamics. But is an additional feeding mode needed for understanding and predicting salmonid population responses to streamflow and other environmental factors? We addressed this question by applying two versions of the individual-based model inSTREAM to a field experiment in which streamflow was varied in experimental units that each contained a stream pool and the adjacent upstream riffle. The two model versions differed only in the feeding options available to fish. Both versions of inSTREAM included drift feeding; one also included a search feeding mode to represent feeding in which food availability is largely independent of streamflow, such as feeding from the benthos, or feeding from the water column or the water’s surface in low water velocities. We compared the abilities of the two model versions to fit the observed distributions of growth by individual rainbow trout (Oncorhynchus mykiss) in the field experiment. The version giving fish the daily choice between drift or search feeding better fit observations than the version in which fish fed only on drift. Values for drift and search food availability from calibration to the individual mass changes of fish in experimental units with unaltered streamflow yielded realistic distributions of individual growth when applied to experimental units in which streamflow was reduced by 80 %. These results correspond with empirical studies that show search feeding can be an important alternative to drift feeding for salmonids in some settings, and indicate that relatively simple formulations of both processes in individual-based population models can be useful in predicting the effects of environmental alterations on fish populations.     

Incorporating invertebrate predators into theory regarding the timing of invertebrate drift

Theory concerning the timing of lotic invertebrate drift suggests that daytime-feeding fish cause invertebrates to restrict their drift behavior to the nighttime. However, there is growing evidence that the nighttime foraging of invertebrate predators also contributes to the nocturnal timing of drift, though it is unclear whether the nocturnal behavior of invertebrate predators is innate or proximately caused by fish. In two experiments, one conducted in a fish-bearing stream and a second in a fishless stream, we compared the drift patterns of Baetidae (Ephemeroptera) from channels with and without benthic invertebrate predators. We tested whether invertebrate predators affect the timing of drift, either as a proximate cause of nocturnal drift in the fishless stream (diel periodicity) or as a proximate cause of a pre-dawn peak in drift in the fish-bearing stream (nocturnal periodicity). In the fish-bearing stream experiment, a pre-dawn increase of baetid drift occurred independently of invertebrate predators, indicating that invertebrate predators were not the proximate cause of nocturnal periodicity in the stream. In the fishless stream experiment, invertebrate predators caused more baetid drift at night than during the day, indicating that invertebrate predators caused the nocturnal drift pattern we observed in the stream, and that invertebrate predators can influence drift timing independently of fish. Therefore, we suggest that both visually feeding fish and nocturnally foraging benthic predators, when present, affect the timing of invertebrate drift; visually feeding fish by reducing daytime drift, and benthic predators by increasing nighttime drift.     

Short-term study testing the resilience of an estuarine bivalve to macroalgal mats

Increased mortality of juvenile Atlantic salmon (Salmo salar L.), related to lowered levels of stored energy following the loss of ice cover during winter, has been observed after hydropower development in the subarctic River Alta, northern Norway. Drift samples were compared to examine if drift densities, and thus drift prey availabilities for juvenile salmon, were lower in the ice-free than the ice-covered area. In addition, juvenile salmon stomach contents were compared to benthos and drift in the ice-free area to examine salmon winter feeding habitat. Zooplankton, originating from the reservoir, dominated drift at the ice-free site but had lower densities at the downstream ice-covered site. Excluding zooplankton, Chironomidae comprised most of the remaining drift at both the ice-free and ice-covered site, followed by Ephemeroptera, Plecoptera and Simuliidae. No Trichoptera were found in the drift samples. There was no consistent diel periodicity in drift. Benthos was dominated by Chironomidae, followed by Ephemeroptera, Plecoptera and Trichoptera. Other invertebrates occurred in low numbers. Juvenile salmon demonstrated size-selective feeding and fed mainly on Ephemeroptera, followed by Trichoptera and Plecoptera. No zooplankton and few Chironomidae were found in the stomach samples. Stomach content was more similar to benthos than to drift, indicating a larger extent of benthic than drift feeding. No evidence was found for the hypothesis that lack of ice cover reduced the invertebrate drift or caused diel periodicity in the drift. Differences in drift between areas with and without ice could not account for the observed differences in mortality of juvenile salmon during the winter in these areas.     

Substrate selection and seasonal variation in densities of invertebrates in stream pools of a tropical river

Spatial distribution and seasonal variation in densities of the invertebrates were investigated for a year in three stream pools of a South Indian river. The effects of season, substrate type and water depth on the distribution were analyzed. Substrate type and season influenced the invertebrate distribution the most. Leaf packs harboured most of the organisms followed by macro-algal substrate and sand. The lowest densities were observed on rocky substrates and in the water column. Rocky substrates in shallow water supported higher densities of total invertebrates than deeper areas. Chironomid larvae dominated all benthic substrates throughout the year. Of the 19 invertebrate taxa studied, 6 showed no seasonality in densities, and most of the rest showed their highest densities in the pre-monsoon period and lowest in the SW monsoon or post-monsoon periods. However, in two of the three pools, the densities of total invertebrates were highest during the post-monsoon period with secondary peaks in the pre-monsoon period.     

Invertebrate drift lost to the sea during low flow conditions in a small coastal stream in Western Canada

The invertebrate drift leaving the lower end of a small coastal stream on the east coast of Vancouver Island was recorded during low flow conditions. 20,156 animals, weighing 0.96 g (dry weight), were collected in drift nets over a 5 day period in early summer. High and low drifting taxa are listed. Considerable daily variation in total captures occurred and was attributed to a few dominant taxa (primarily harpacticoid copepods, mites and chironomid larvae) exhibiting atypical drift patterns. 0.004% of the stream's invertebrate standing crop was estimated to be in the water column at any instant in time. The possible use of the outgoing animals as food for juvenile salmon in the estuary is discussed.     

Temporal consistency in the long-term spatial distribution of macroinvertebrate drift along a stream reach

Previous studies of the spatial pattern of stream invertebrate drift have focused on spatial variation at microhabitat scales or landscape scales, or temporal variation over diel or seasonal scales. None have examined consistency in spatial variation over longer time scales (>1 year). This study examined invertebrate drift density and composition at fixed locations (terminal ends of 10 riffles) each month at day and night along a 1 km reach of a 2nd order stream over a period of nearly 2 years. Consistent differences in the density of macroinvertebrate drift between riffles over 2 years were observed. The only habitat characteristic observed to be related to invertebrate drift density was the length and size of riffles above sampling sites, with larger and longer riffles producing the highest drift densities. Consistent differences in the supply of drifting macroinvertebrates along a stream reach may have implications for the supply of colonists to substrate patches and the profitability of feeding positions for drift-feeding fish and other predators. Handling editor: D. Dudgeon     

Spatial and temporal variability of macroinvertebrates in spawning and non-spawning habitats during a salmon run in Southeast Alaska

Spawning salmon create patches of disturbance through redd digging which can reduce macroinvertebrate abundance and biomass in spawning habitat. We asked whether displaced invertebrates use non-spawning habitats as refugia in streams. Our study explored how the spatial and temporal distribution of macroinvertebrates changed during a pink salmon (Oncorhynchus gorbuscha) spawning run and compared macroinvertebrates in spawning (riffle) and non-spawning (refugia) habitats in an Alaskan stream. Potential refugia included: pools, stream margins and the hyporheic zone, and we also sampled invertebrate drift. We predicted that macroinvertebrates would decline in riffles and increase in drift and refugia habitats during salmon spawning. We observed a reduction in the density, biomass and taxonomic richness of macroinvertebrates in riffles during spawning. There was no change in pool and margin invertebrate communities, except insect biomass declined in pools during the spawning period. Macroinvertebrate density was greater in the hyporheic zone and macroinvertebrate density and richness increased in the drift during spawning. We observed significant invertebrate declines within spawning habitat; however in non-spawning habitat, there were less pronounced changes in invertebrate density and richness. The results observed may be due to spawning-related disturbances, insect phenology, or other variables. We propose that certain in-stream habitats could be important for the persistence of macroinvertebrates during salmon spawning in a Southeast Alaskan stream.