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.     

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.     

The impact of physical disturbance on stream communities: lessons from road culverts

This study examined the impact of physical disturbance from long-established road culverts on stream macroinvertebrate communities. Three streams within a 6 km section of highway on the Avalon Peninsula, Newfoundland, Canada, were sampled. Streams had the entire upstream watershed and at least 100 m downstream of the road with natural boreal forest/barren vegetation and all had, within the sampled reaches, similar physical streambed characteristics. The fauna on stones from riffles was sampled at two upstream and three downstream sites, i.e., from 50 m above to about 100 m below the road in each stream. A total of 33 taxa were identified among the streams, with differences limited to a few rare taxa. The sample site communities did not significantly differ from each other with respect to the taxa present. Total macroinvertebrate abundance by site, for combined data of all streams, indicated the site at the exit of culvert plunge pool (site 3) had significantly elevated abundances. Analysis of individual taxa showed this was primarily due to very high numbers of Simulium spp. The other most notable changes were a decrease in numbers of Hydropsyche spp. and Elmidae below the road. The abundances of the remaining taxa were more variable among all sites. The study indicated that long-standing point source physical disturbance primarily impacted taxa abundance rather than community present/absent data, which will recolonize the disturbed zone by downstream drift. The differences in abundance are probably the result of the cleaning of substrate by abrasion, movement of substrate and reduction of detritus during each spate. Handling editor: D. Dudgeon     

Do pools impede drift dispersal by stream insects?

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Disturbance history influences the distribution of stream invertebrates by altering microhabitat parameters: a field experiment

1. We investigated the effects of local disturbance history and several biotic and abiotic habitat parameters on the microdistribution of benthic invertebrates after an experimental disturbance in a flood‐prone German stream. 2. Bed movement patterns during a moderate flood were simulated by scouring and filling stream bed patches (area 0.49 m²) to a depth of 15–20 cm. Invertebrates were investigated using ceramic tiles as standardized substrata. After 1, 8, 22, 29, 36 and 50 days, we sampled one tile from each of 16 replicates of three bed stability treatments (scour, fill and stable controls). For each tile, we also determined water depth, near‐bed current velocity, the grain size of the substratum beneath the tile, epilithic algal biomass and standing stock of particulate organic matter (POM). 3. Shortly after disturbance, total invertebrate density, taxon richness and density of the common taxa Baetis spp. and Chironomidae were highest in stable patches. Several weeks after disturbance, by contrast, Baetis spp. and Hydropsychidae were most common in fill and Leuctra spp. in scour patches. The black fly Simulium spp. was most abundant in fill patches from the first day onwards. Community evenness was highest in scour patches during the entire study. 4. Local disturbance history also influenced algal biomass and POM standing stock at the beginning of the experiment, and water depth, current velocity and substratum grain size throughout the experiment. Scouring mainly exposed finer substrata and caused local depressions in the stream bed characterized by slower near‐bed current velocity. Algal biomass was higher in stable and scour patches and POM was highest in scour patches. In turn, all five common invertebrate taxa were frequently correlated with one or two of these habitat parameters. 5. Our results suggest that several ‘direct’ initial effects of local disturbance history on the invertebrates were subsequently replaced by ‘indirect’ effects of disturbance history (via disturbance‐induced changes in habitat parameters such as current velocity or food).     

Meiofauna constitute a considerable portion of invertebrate drift among moss-rich patches within a karst hydrosystem

Aiming to establish the most frequent invertebrate taxa in drift at the small spatial scale within a moss-rich karst tufa-precipitating hydrosystem, we sampled drift among microhabitats differing in substratum type and flow conditions along a tufa barrier-cascading lotic reach. Additionally, we addressed the question of the contribution and the potential significance of meiofauna within the overall invertebrate drift at the small spatial scale. During the study period, a total of 60 invertebrate taxa were recorded in the drift. Six of these taxa belonged to the annelid/arthropod meiofauna and they represented 35% of total drift density. Macroinvertebrates found in drift were represented mainly by larval insects. The composition of the most abundant taxa in total drift was as follows: Alona spp. (Cladocera 26.7%), Riolus spp. (Coleoptera: Elmidae 13.2%), Simulium spp. (Diptera: Simuliidae 12.2%), Enchytraeidae (Oligochaeta 10.4%), Hydrachnidia (6.3%), Orthocladinae (Diptera: Chironomidae 3.9%) and Naididae (Oligochaeta 3.6%). Faunal drift densities and amounts of transported particulate matter (PM) were highest at the fast-flowing sites located at the barriers and lowest at the slow-flowing sites within pools. Similarly to the seasonal amounts of transported PM, faunal drift was lowest in winter, and peaked in autumn and in late spring/early summer. Correlation between flow velocity and PM-faunal drift densities suggested a significant effect of the dislodged PM, though a minor influence of discharge and flow velocity on faunal drift. We suggest that the small-scale habitat heterogeneity and the respective feeding and refugial strategies of the fauna, as well as faunal passive dislodgement initiated by the shear forces of the flow were the most important drivers of observed drift patterns.     

Response of benthic invertebrates to natural versus experimental disturbance in a Swiss prealpine river

1. The crucial point of disturbance experiments in streams is the extent to which they can simulate a natural spate. Ideally, disturbance experiments should proceed side by side with a phenomenological study to allow a direct comparison. In the present study conducted in a prealpine Swiss river, the River Necker, fortuitous events made such a comparison possible. 2. In summer 1994, we took Surber samples one day before and on several sampling dates after a major flood (recurrence interval ≈ 5 years), which was followed by a long period of uniform discharge in a river characterized by frequent spates. Beginning 19 days after this flood, patches of the stream bed (≈ 9 m²) were physically disturbed by kicking and raking. 3. The degrees of reduction in the total number of individuals and the dominant taxa were similar after both types of disturbance, as were the recolonization patterns of Rhithrogena spp., Leuctra spp. and Hydracarina. Chironomidae, Baetis spp., Simuliidae, Pentaneurini and Corynoneura/Thienemanniella spp. showed a distinct lag phase after the flood before recolonization began, whereas there was no such lag phase after the experiment. Therefore, the time needed to recover to pre-flood densities was longer for these taxa. Nevertheless, recolonization rates and patterns after the lag phase were similar to those after the experimental disturbance. 4. Size-class measurements indicated that recruitment from egg hatching may have been more important after the flood than after the experimental disturbance for Rhithrogena spp., but not for Chironomidae, Baetis spp., Simuliidae, Pentaneurini and Leuctra spp. Invertebrate drift was probably the most important pathway of recolonization after both types of disturbance. 5. Our experiment allowed a realistic simulation of several important effects of the large flood on the invertebrate community. Smaller spates that induce substratum movement at a spatial scale similar to our experimental plots are much more common than large floods in the River Necker. For these spates, our experiment should provide an even more realistic simulation of natural disturbance.     

Downstream drift of invertebrates in a river in southern Ghana

The results of quantitative sampling of the drifting invertebrates from a riffle area in the upper reaches of the Pawmpawm River in the forest of Ghana are presented. The observations were made once a month during 14 months, April 1970 to May 1971, and also on a daily basis for 3 weeks. Changes in numbers and composition of the animals drifting were examined in relation to the volume of water flowing and the current; there appeared to be two distinct drift phenomena, ‘true’ and ‘background’ drift. The animals belonging to each of these two components are discussed in relation to their response to light, to diurnal fluctuations in total drift and to the phases of the moon. There was a peak drift activity after sunset in those animals which are affected by light, e.g. Centroptilum, Austrocaenis, Chironomidae, Simulium. It is concluded that drift may be simply a function of the activity patterns of the insects.     

Macroinvertebrate drift in a Rocky Mountain stream

An extensive series of drift collections from a Rocky Mountain stream was used to investigate quantitative patterns in the taxonomic composition of drift throughout spring, summer and fall for 1975–1978. Drift was estimated by drift rate, the number of organisms drifting past a point per 24 h; and by drift density, the numbers of organisms collected per 100 m³ of water sampled.Drift densities were up to ten times greater by night than by day, and 24 h drift densities for the total fauna approached 2000 per 100 m³ in June–July, declining to <500 by autumn. Ephemeroptera, and especially Baetis, dominated the drift. Drift rates were greatest in late spring, around 10⁶ per 24 h, which are among the highest values reported for small trout streams. Drift rates declined to <10⁵ during the summer, and shifts in the taxonomic composition are described.Multiple regression analysis of the relationship between drift rate and density, and the independent variables discharge, benthic density and temperature, showed that discharge typically was a significant predictor of 24 h drift rate, usually the best single predictor. In contrast, 24 h drift density most frequently was independent of discharge, indicating that this measure tends to correct for seasonal variation in discharge, as suggested in the literature. However, this was not invariably true. Drift density significantly correlated with benthic density in five of eight taxa inspected, thus seasonal declines in the benthos probably accounted for parallel declines in drift density.     

Seasonal fluctuations of macroinvertebrate drift in a South Carolina Piedmont stream

The seasonal fluctuations of larval macroinvertebrate drift, exuvial drift and larval benthic density were quantitatively examined over a 1-year period in a fourth order, spring-fed stream in the Piedmont area of South Carolina. The drift was dominated by the mayfly Baetis spp. and by two species of blackfly (Prosimulium mixtum and Simulium jenningsi). Peak drift densities were noted during early spring and especially late summer. Strong correlations were noted between larval drift densities and exuvial drift, indicating a relationship between drift and seasonal growth and emergence patterns. Seasonal trends in drift and benthic densities, though less strongly correlated, were also generally similar.     

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.     

Benthic macroinvertebrate distributions in the riffle-pool communities of two east Texas streams

The benthic macroinvertebrate riffle-pool communities of two east Texas streams were sampled monthly for a period of one year. In contrast to previous studies in primarily upland areas, pools in Alazan Creek and Bernaldo Bayou contained significantly higher densities and biomass, as well as a significantly higher diversity and number of taxa. The majority of taxa collected could be characterized as pool adapted organisms, with the Diptera, Ephemeroptera, Coleoptera, and Odonata having greater numbers in pools. Perlesta, Baetis, and Cheumatopsyche were the only genera to have significant numbers in riffles at both sampling sites. A greater number of taxa were restricted to pools at both sampling locations, and although an unstable, sandy substrate was a factor in limiting colonization of riffles, riffles were not as productive as pools even in the presence of suitable substrate conditions. A cluster analysis based on similarity indices indicated that like habitats between streams were more similar to each other, than were adjacent riffle-pool complexes within streams.     

Aquatic insect drift through a final-cut strip mine pit,with emphasis on drift distances

Drift samples were taken upstream and at the outlet of a final-cut strip mine pit through which a portion of the Tongue River, Wyoming, USA, was rerouted. All prominent rheophilic insect species (except cased Trichoptera) were found capable of drifting the length of the pit (> 500 m). However, drift densities leaving the pit were reduced in comparison to those entering it. Pit inhabitants such as burrowing mayflies, Chaoborus flavicans, and several species of Chironomidae were also collected at the pit outlet. Under the physical conditions created by rerouting the river through a strip mine pit, the observed maximum behavioral and background drift distances (> 500 m) were significantly greater than those reported in the past (< 100 m).     

The drifting of invertebrates and particulate organic matter in an Austrian mountain brook

1. The drift of organisms and particulate organic matter in a calcareous mountain brook near Lunz, Lower Austria, was investigated for 1 year. Five sets of three drift nets (sampling at different water depths) were distributed along a cross-section of the brook. 2. From February to March 1989, samples were taken at intervals of 2–7 days over 24-h periods, at the end of which the drift nets were emptied. From April 1989 to March 1990, sampling was carried out at monthly intervals and the nets emptied every 3h for 1 full day. 3. Of 71 810 organisms caught, the most abundant taxa were Diptera (46.6%; mainly Chironomidae and Simuliidae), Ephemeroptera (24.0%; dominated by Baetis spp.) and Plecoptera (16.1%). 4. The drift density (mean ± 95% CL) was significantly higher (P<0.05) in spring and summer (2.50 ± 0.32 specimens m^?3) than in autumn and winter (2.01 ± 0,22 specimens m^?3). In terms of biomass (wet weight), drift density was also significantly higher (P < 0.001) in spring and summer (2.50 ± 0.48 mgm^?3) than during the rest of the year (1.04±0.12mgm ^?3). 5. Total drift rates through a cross-section of the brook ranged from 17366 organisms per day at low discharge (water depth = 10cm) to 955152 individuals per day at a water depth of 50cm in autumn and winter; in spring and summer the corresponding values were 21600 and 1188000 specimens per day. 6. Ephemeroptera, Plecoptera, Coleoptera and Simuliidae were most abundant in night samples, whereas Hydracarina, Trichoptera and Chironomidae drifted mainly during daylight hours. 7. Drift density (dry weight) of particulate organic matter was positively correlated with discharge and ranged from 2.16mgm^?3 at a water depth of 10cm to 17.39mgm^?3 at 50cm.     

Fish predation affects the structure of a benthic community

1. We conducted an experimental study of predation by benthivorous fish on a natural community of stream invertebrates using a reach‐scale approach. Over a 2‐year period (experimental phase), the benthic invertebrate community of a stretch containing two species of benthivorous fish was compared with a fishless stretch. Thereafter, all fish were removed and benthic community structure was analysed again to account for natural differences between the two stretches (reference phase). 2. Benthivorous fish at the moderate densities investigated did not affect total benthic biomass or density, but did alter species composition. In addition, the fish effect differed between pool and riffle habitats, with larger effects in the pools indicating a habitat‐specific predation effect. In the reference phase, when all fish were removed from the stream, the difference between the two stretches was reduced. 3. The benthivorous fish reduced the densities of four taxa (Pisidium sp., Dugesia gonocephala, Gammarus pulex, Limoniidae), representing 29% of total biomass. It is possible that density reductions of other species were masked by prey migration despite the relatively large spatial scale. Indeed, higher drift activity in the upstream fishless stretch could have increased the density of Baetis rhodani in the fish stretch, as indicated by the results of a drift model. 4. Our results provide insights into stream food web ecology because fish predation showed effects even in a natural system where habitat complexity was high, environmental factors were highly variable and many predator and prey species interacted and because benthivorous fish were the focus, whereas the majority of previous predation experiments in streams have used drift‐feeding trout.     

Size-dependent drift responses of mayflies to experimental hydrologic variation: active predator avoidance or passive hydrodynamic displacement?

Summary Larger nymphs within aquatic insect taxa have been frequently observed to be transported down-stream in the stream drift only at night. Others have hypothesized this pattern results primarily from large nymphs' behavioural avoidance of entering drift during daylight, when size-selective, visually-feeding fish predators are most active. This hypothesis assumes that animals can actively control their entry into the drift, which may not be the case under all flow conditions. We experimentally induced streamflow increases and decreases in adjacent riffles in a hydrologically-stable stream during the daytime to examine whether changes in diel patterns of drift abundance and size-distribution of mayflies were consistent with the hypothesis of active avoidance of diurnal drift. We assessed the likelihood of active vs. passive mechanisms of diurnal drift entry and transport for four taxa that differ with respect to body size, morpho-behavioural attributes, microhabitat use, and general propensity to drift. In each of three seasons, diurnal and nocturnal drift samples were collected in three riffles over two diel cycles. Background drift patterns were established on the first day (no flow manipulation). Six h before sunset on the second day, flow was experimentally increased in one riffle, decreased in the second, and not altered in the third (control). Between-day differences in diurnal and nocturnal drift rate and size composition were then compared among the treatment and reference riffles. Responses of two taxa were consistent with active control over drift entry, transport, or both. For Baetis spp., drift-prone mayflies typically preyed upon by fish, diurnal drift rates immediately increased following both flow reduction and flow elevation in all seasons, but only small individuals comprised the drift. Drift by large individuals was delayed until nighttime. Epeorus longimanus also exhibited significant increases in drift rates following flow reduction and elevation, but responses of this large-bodied species were restricted to nighttime. Drift responses for these two taxa were largely independent of direction of hydrologic change, thus indicating a strong behavioural control over drift. By contrast, numbers and sizes of drifting Paraleptophlebia heteronea and Ephemerella infrequens depended strongly on direction of flow change. Drift rates for both species generally declined after flow reduction and increased after flow elevation. Moreover, after flow elevation, larger individuals often drifted diurnally, a finding consistent with expectations under a passive hydrodynamic model. These experiments indicate that size-dependent mayfly drift reflects not only presumed risk from visual fish predators, but also functional attributes of species such as morphology, behaviour, and microhabitat affiliation, which influence aspects of drift entry and transport under variable hydrologic conditions.     

Spatial and temporal patterns in the fish assemblages of individual pools in a midwestern stream (U.S.A.)

Synopsis The composition and consistency of fish assemblages in 14 adjacent pools (6–120 m long) of a clear-water, limestone and gravel creek in midwestern U.S.A. were quantified in eight snorkeling surveys over 19 months, to establish a baseline of natural variation in the system at this scale. The fauna of the stream was dominated numerically by minnows (Cyprinidae), sunfish and black bass (Centrarchidae), and topminnows (Fundulidae). The pool fish fauna of the total 1 km reach (including all 14 pools) was highly consistent throughout the study, despite two major floods. Assemblages in individual pools generally were consistent, but there was more variation within pools than at the scale of the entire reach. Throughout the study, most individual pools remained within discrete subsets of the total occupied multivariate space in a principal components analysis based on fish species abundances. Sunfishes (Lepomis spp.) and bass (Micropterus spp.) were more consistent in their distribution among pools than were minnows (Cyprinidae) or a topminnow (Fundulus). There were 25 significant correlations in occurrence of species pairs among stream pools, out of 91 possible comparisons of the 14 most abundant taxa in the reach. Many pools contained assemblages either dominated by large centrarchids or by abundant cyprinids and juvenile centrarchids, but intermediate assemblages also were observed. The dynamics of distribution of fish species and fish assemblages among individual stream pools are likely influenced by a combination of species-specific behaviors and habitat selection, predator constraints on use of individual pools by small fishes, riffles as size-selective barriers to fish movements between pools, dispersal of young-of-the-year fishes, and abiotic phenomena like floods. Individual stream pools appear to be discrete habitat units for fishes, and do represent an appropriate scale for biologically meaningful studies of fish assemblages or their effects on streams.Department of Zoology, University of Oklahoma     

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.     

Response of endangered Poeciliopsis occidentalis sonoriensis in the Río Yaqui drainage,Arizona, to introduced Gambusia affinis

Synopsis Potential coexistence of the native Yaqui topminnow, Poeciliopsis occidentalis sonoriensis, with introduced mosquitofish, Gambusia affinis, was examined in spring pools and streams in San Bernardino National Wildlife Refuge, southeastern Arizona. Poeciliopsis never exceeded 12% of total poeciliid numbers in a spring pool where Gambusia was present. Body size, fecundity and reproductive effort of Poeciliopsis were significantly higher in this pool than in similar spring pools where Gambusia was absent. Where Poeciliopsis and Gambusia were syntopic in a stream, numbers of Poeciliopsis declined over 1.7 years, until none were collected. Poeciliopsis then increased to > 60% of total poeciliid numbers following flash flooding. Persistence of Poeciliopsis with Gambusia in the spring pool appeared to be a result of compensatory increase in reproductive output, while in the stream it was associated with recurrent flash flooding and a uniform temperature springhead which provided refuge.     

Comparisons of benthic invertebrates between riffles and pools

Benthic species assemblages in upstream and downstream ends of riffles and in pools were investigated seasonally in the first five orders of an alluvial gravel stream with distinct pool and riffle channel form. Riffles comprised < 10% of stream area and were separated by pools with extensive bedrock substrate (ca. 15–85% of total surface area) which was scoured during periodic high flow. Virtually all taxa were more abundant in riffles than in pools, except chironomids which were more equally distributed. Inconsistent results were obtained for upstream-downstream comparisons within riffles. Intermittent headwater reaches (orders 1 & 2) which supported half as many taxa retained this pattern during periods of flow, although riffles at these sites were dry from mid-June to mid-November. Pools which contained more gravel, indicating less disturbance during high flow, had a richer assemblage of benthic species than other pools. Many invertebrates in pools may have been there as a result of drift from their preferred riffle habitats, but the presence of gravel in the pools indicates less intense flow disturbance during floods, provides protection from the mild scouring that does occur during floods, and provides refugia from predators.