Local disturbance history and habitat parameters influence the microdistribution of stream invertebrates

1. We investigated the effects of local disturbance history and habitat parameters (abiotic and biotic) on the microdistribution of benthic invertebrates during several floods in two streams, the Schmiedlaine in Germany (four events) and the Kye Burn in New Zealand (two events). 2. Bed movement patterns were quantified using metal‐link scour chains. Before and after each flood, quantitative invertebrate samples were taken from replicate bed patches that had experienced sediment scour, fill or remained stable. 3. Patterns of invertebrate density in the different bed stability types (i.e. scour, fill, stable) varied between floods, sampling dates and streams, but invertebrate density was highest in stable patches in >50% of all the patch type effects detected and lowest in fill patches in 75% of all detected effects. Stable bed patches acted as a refugium for Liponeura spp. and Leuctra spp. in the Schmiedlaine and for Hydracarina and Deleatidium spp. in the Kye Burn. 4. Averaged across both streams, only near‐bed current velocity was correlated with invertebrate distribution on the streambed more often than disturbance history. In the Kye Burn, disturbance history and water depth were the most influential habitat parameters. 5. Our results suggest that a thorough understanding of the microdistribution of benthic invertebrates requires knowledge of disturbance history, as well as more readily measured habitat parameters such as current velocity or water depth.     

Exotic crayfish in a brown water stream: effects on juvenile trout, invertebrates and algae

SUMMARY 1. The impact of the introduced omnivorous signal crayfish (Pacifastacus leniusculus) on trout fry, macroinvertebrates and algae was evaluated in a brown water stream in southern Sweden using in situ enclosures. We also examined the gut content of all surviving crayfish in the enclosures. Two crayfish densities in addition to a control without crayfish were used in replicate enclosures (1.26 m²) in a 1‐month experiment. Additionally, 20 trout fry (Salmo trutta) were stocked in each enclosure to assess the effects of crayfish on trout survival and growth. 2. Detritus was the most common food item in crayfish guts. Animal fragments were also frequent while algae and macrophytes were scarcer. Crayfish exuviae were found in crayfish guts, but the frequency of cannibalism was low. 3. Trout survival in enclosures was positively related to water velocity but was unaffected by crayfish. 4. Total invertebrate biomass and taxon richness were lower in crayfish treatments. The biomass of all predatory invertebrate taxa was reduced but only three of six non‐predatory taxa were reduced in the crayfish treatments. 5. Epiphytic algal biomass (measured as chlorophyll a, on plastic strips) was not related to crayfish density, whereas the biomass of epilithic algae (measured as chlorophyll a) was enhanced by high water velocity and high crayfish density. The latter was possibly mediated via improved light and nutrient conditions, as active crayfish re‐suspend and/or remove detritus and senescent algal cells during periods of low water velocity. 6. We conclude that the introduced signal crayfish may affect stream communities directly and indirectly. Invaded communities will have reduced macroinvertebrate taxon richness and the signal crayfish will replace vulnerable invertebrate predators such as leeches. In streams that transport large amounts of sediment or organic matter, a high density of crayfish is likely to enhance benthic algal production through physical activity rather than via trophic effects.     

Selective foraging on terrestrial invertebrates by rainbow trout in a forested headwater stream in northern Japan

The important contribution of terrestrial invertebrates to the energy budget of drift-foraging fishes has been well documented in many forested headwater streams. However, relatively little attention has been focused on the behavioral mechanisms behind such intensive exploitation. We tested for the hypothesis that active prey selection by fishes would be an important determinant of terrestrial invertebrates contribution to fish diets in a forested headwater stream in northern Japan. Rainbow trout, Oncorhynchus mykiss, were estimated to consume 57.12 mg m^–2 day^–1 (dry mass) terrestrial invertebrates, 77% of their total input (73.89 mg m^–2 day^–1), there being high selectivity for the former from stream drift. Both the falling input and drift of terrestrial invertebrates peaked at around dusk, decreasing dramatically toward midnight. In contrast, both aquatic insect adults and benthic invertebrates showed pronounced nocturnal drift. Because the prey consumption rates of rainbow trout were high at dawn and dusk, decreasing around midnight, the greater contribution of terrestrial invertebrates to trout diet was regarded as being partly influenced by the difference in diel periodicity of availability among prey categories. In addition, selectivity also depended upon differences in individual prey size among aquatic insect adults, and benthic and terrestrial invertebrates, the last category being largest in both the stream drift and the trout diets. We concluded that differences in both the timing of supplies and prey size among the three prey categories were the primary factors behind the selective foraging on terrestrial invertebrates by rainbow trout.     

Determining the density dependence of immigration and emigration of benthic stream invertebrates: theoretical considerations

Simple mathematical models are formulated to describe density independent and density dependent dispersal. These models clarify hypotheses of density dependence and may be manipulated easily to suit particular applications. The models demonstrate that the initial composition of a species aggregate must be controlled before valid conclusions can be drawn about the density dependency of the aggregate's dispersal. Stochastic models of emigration are derived to assess the power of particular experimental designs and statistical techniques to discriminate a known form of density dependent emigration. Contribution No. 369, Great Lakes Research Division, University of Michigan Contribution No. 369, Great Lakes Research Division, University of Michigan     

An experimental test of the effects of food resources and hydraulic refuge on patch colonization by stream macroinvertebrates during spates

1. The passive or active movement of organisms between habitat patches plays important roles in achieving ecosystem resilience to disturbance and long-term control of population levels. However, causal mechanisms of disturbance-induced movements of mobile biota across heterogeneous habitat patches at a relatively short time-scale are little understood. 2. We experimentally tested the effects of food resource values on macroinvertebrate colonization of hydraulic refugia from spates in a second-order creek. Experimental cages were colonized by macroinvertebrates with combinations of resource types (natural or polyester leaves), and extent of exposure to stream flow (exposed to or sheltered from current); one half of each set was collected before and after a spate. This experiment was repeated over three spates of varying magnitude and seasonal contexts. 3. Pre-spate colonization was consistently greater for the cages with natural leaves relative to artificial leaves regardless of the extent of flow exposure. Two autumn spates with relatively low and stable antecedent flow conditions caused large movements of organic matter and macroinvertebrates across the stream, showing community-level accumulations into hydraulically sheltered patches independent of food treatment. The smallest spate with high and variable antecedent flows during winter resulted in negligible responses, which we interpret to be a result of depletion of easily transportable organic matter and organisms. 4. Two detritivorous taxa, the mayfly Paraleptophlebia spp. and stonefly Despaxia augusta (Banks) were the most responsive to autumn spates, and had disproportionately higher colonization rates of cages when provided with natural leaves during the largest autumn spate. Preferential settlement in food-enriched hydraulic refugia was attributable to taxon-specific mobility related to efficient acquisition of detritus resource, whose availability varies spatially and temporally. 5. Our findings suggest (1) detritivorous macroinvertebrate colonization of hydraulic refugia can be influenced by hydraulic controls as well as food resource value, and (2) pre-spate environmental conditions in terms of resource distribution and availability may pre-condition organisms' susceptibility to spates and also affect refugium usage, at least in food-limited, detritus-based stream systems.     

The relative contributions of refugium types to the persistence of benthic invertebrates in a seasonal snowmelt flood

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Influences of habitat complexity on the diversity and abundance of epiphytic invertebrates on plants

SUMMARY 1. The compound influence of habitat complexity and patch size on stream invertebrate assemblages associated with submerged macrophytes was investigated through field sampling of two natural macrophyte species with contrasting leaf morphologies (complex, Ranunculus yezoensis; simple, Sparganium emersum) and an experiment with two artificial plants with different levels of morphological complexity. 2. The artificial plant experiment was designed to separate the effects of habitat area (patch size) and habitat complexity, thus enabling a more rigorous assessment of complexity per se than in previous studies where only a single patch size was used. Simple and complex artificial plants were established with five different patch sizes corresponding to the range found in natural plants. 3. Invertebrates occurred on both complex and simple forms of natural and artificial plants at similar abundances with dipterans and ephemeropterans being predominant. Taxon richness was higher on structurally complex Ranunculus than on simple Sparganium and was similarly higher on the complex artificial plant than on the simple one, over the entire range of habitat patch sizes. Thus, architectural complexity affected the taxon richness of epiphytic invertebrates, independently of habitat scale. 4. On the natural plants there was no difference in the abundance (both number of individuals and biomass) of invertebrates between simple and complex forms, while on artificial plants more invertebrates occurred on complex than on simple forms. The amount of particulate organic matter, >225 μm (POM) and chlorophyll a showed mixed patterns on natural and artificial plants, suggesting that the availability of these resources is not an overriding proximate factor controlling invertebrate abundance on plants. The difficulty of extrapolating from experimental results involving use of artificial plants is discussed, especially when considering the relationship between habitat structure and the occurrence of epiphytic invertebrates on natural plants.     

Spatial and temporal occurrence of invertebrates in a chalk stream,Berkshire, England

The invertebrate assemblages of the River Lambourn and its tributary, the Winterbourne stream, were investigated as part of a broad ecological study prior to water resource development in the catchment. The longitudinal distribution of the invertebrate fauna within the Winterbourne stream was examined in detail and supporting data for the R. Lambourn revealed that the upstream intermittent section of each water course had an invertebrate assemblage which was distinct from the fauna downstream where flow was permanent. Within the perennial zone taxon richness tended to increase downstream.A one-year programme of monthly quantitative sampling on five distinct biotopes (Ranunculus, Berula, Callitriche, gravel and silt) at two contrasted sites (shaded and unshaded) was undertaken in the lower perennial of the R. Lambourn below the confluence with the Winterbourne. On each study site, the taxon richness, numerical abundance and biomass of invertebrates were significantly higher on the three macrophytes than on gravel and silt. There were also statistically significant seasonal changes in the biomass of invertebrates on each site. Estimates of total biomass of invertebrates per site revealed a late spring peak (May/June) and a late autumn (November/December) peak, neither of which coincided with the summer peak biomass of macrophytes. This suggested that, in the lower perennial, the level of food resources (epiphytic algae in late spring and fallen tree leaves plus decaying macrophytes in autumn) rather than available habitat, were a major influence on population levels for most of the year.Quantitative sampling of each biotope on each site continued in June and December only for a period of 8 years (1971–78), during which time the river experienced a minor drought (1973) and a severe drought (1976). The Chironomidae showed a strong positive response to conditions of low flow in June of 1973 and 1976, when exceptionally high densities of larvae were recorded. Although further between-year differences were recorded in several other major components of the invertebrate fauna, which may have been related to conditions of low discharge, they did not have long-term consequences for the fauna in the lower perennial of the R. Lambourn.In contrast, in the upper perennial, the prolonged drought of 1976 resulted in the loss of wetted perimeter and macrophytes, coupled with deposition of silt, all of which had deleterious effects on the invertebrate assemblages. The following year, despite a normal discharge regime and the removal of silt, some submerged macrophytes were slow to return to their pre-drought condition. In the intermittent section, the drought of 1976 resulted in the total loss of flow for a 16 month period instead of a more typical four month period of water loss.Thus, chalk streams encompass within a single system both stable and extreme hydrological regimes which offer unique opportunities for investigating the processes of community formation and maintenance.     

Microform bed clusters: are they preferred habitats for invertebrates in a flood‐prone stream?

SUMMARY 1. Microform bed clusters are bedform microunits in streams. They consist of an obstacle clast (a large surface stone) against which other stones are stacked in a specific manner. Stream ecologists have suggested that these clusters are better flood refugia and more valuable habitats for invertebrates than single surface stones because of their higher stability during high‐flow events and their greater diversity of microhabitats available for colonisation. 2. To test these predictions in the Schmiedlaine, a flood‐prone prealpine stream, we sampled invertebrates on clustered and single stones on a total of seven occasions including before and after a moderate spate, two major floods and a minor spate. We also monitored surface particle stability during the second major flood and the minor spate. 3. Before the second flood (return period 1–2 years), we determined the exact positions of 60 clusters (60 obstacle clasts with 136 associated smaller particles) and 50 single stones in a 250‐m reach. We sampled the fauna on stones in the clusters, on well‐embedded single stones, and on loose single stones. Only six obstacle clasts and one single stone remained after the flood, implying movement of almost the entire surface layer. Therefore, we sampled the invertebrates on stones in newly formed clusters, on well‐embedded and single loose stones 3 and 11 days after the flood. The minor spate, in contrast to the second flood, moved only one of 495 monitored surface particles. 4. Three days after the flood, invertebrate density on clustered stones and both single stone categories were similar and equalled 34% of the mean pre‐flood density. Eight days later, density had almost doubled. The relatively high survival and rapid recovery suggest that invertebrates found refugia during the flood. However, the very low stability of clusters and single stones implied that surface particles were unimportant as refugia. 5. Total invertebrate density and taxon richness were never higher on clustered than on single stones (regardless of the timing of sampling relative to the last previous high‐flow event). Densities of the seven most common taxa and invertebrate community structure were also generally similar between particle types. We conclude that microform bed clusters cannot be regarded as more valuable invertebrate habitats than single surface stones in the Schmiedlaine.     

Species aggregation: the influence of detritus in a benthic invertebrate community

Species aggregation with detritus was examined in a zoobenthic stream community. Detritus was quantified (dry weight in particle size categories) in contiguously arranged quadrats along four transects within a 40m stream section. Total organism density was found to be positively related to increasing detrital biomass (dry weight) in all transects, although the magnitude of this association was influenced by the detrital particle size distribution in each quadrat. Eliminating certain particle size categories of detritus resulted in substantial changes in the magnitude of association. For example, in one transect, total organism density and total detrital biomass showed no association. However, when one particle size category (greater than 10 mm) was removed a highly significant relationship was found. Total detrital biomass was found to be a useful predictor of organism distribution, however, inordinately high detrital biomass and variance within larger particle size categories can mask a relationship. This result suggests that the use of discrete size categories is necessary when considering species/ detrital relationships.Significant associations were found between the density of particular species and total biomass of detritus. The magnitude, as well as significance, of these associations were found to vary when correlations were calculated with restricted particle size categories of detritus. Species associations were often not consistent between transects, possibly due to local characteristics of detritus or overriding environmental factors. Current velocity was found to be unimportant in determining the detrital biomass held in the substrate. Likewise, few associations were found between current velocity and organism density in three out of four transects. Results of this study suggest that habitat selection by some taxa is influenced by local variation in detrital biomass, as well as detrital particle size composition.     

The role of fungi in the nutrition of stream invertebrates

Dead leaves falling into streams are an important food source for many invertebrates. They are generally made more palatable and more nutritious if they are first colonized by aquatic hyphomycetes and other micro-organisms. At least two mechanisms appear to be responsible for this conditioning effect: microbial production (addition of easily digested microbial compounds to the nutritionally poor leaf substrate), and microbial catalysis (conversion of indigestible leaf substances into digestible subunits by microbial enzymes). Different invertebrate species vary in their ability to take advantage of microbial conditioning. This appears to be influenced by their mobility, the range of their food resources and their ability to overcome defense mechanisms of leaf-colonizing microorganisms.     

Quantifying Abrasion of Stable Substrata in Streams: A New Disturbance Index for Epilithic Biota

Abrasion is a significant disturbing force to which the streambed epilithic biota is exposed during high flow periods. However, abrasion of biota in the field has not been measured, and effects of abrasion on biota are usually inferred. We describe a method that quantifies abrasion of a standardised substrate in the field. This provides an index of the level of abrasion to which flora and fauna are exposed. The method uses autoclaved lightweight aerated concrete (ALC) blocks that are fixed in place on the streambed and subsequently abraded. Two methods were used to quantify abrasion: the change in mass of the block from before to after deployment, and the reduction in cross-sectional area of the corners of the blocks as measured by image analysis. We performed preliminary trials in the laboratory to quantify the amount of work required to remove mass from the blocks. These measurements confirmed that ALC was a consistent substrate with regards to the amount of work required to remove a given mass. The technique was field-tested during a year-long survey at six sites in the Acheron valley, south-eastern Australia. Blocks were deployed for approximately 2 months at a time at each of six sites. The method was able to distinguish average differences in abrasion between trials and sites, but neither of these factors was constant. The variability of abrasion within sites was also high. Results show that the technique in its current form can be used to provide an index of the amount of abrasion to which the epilithic biota is exposed in streams and rivers. An assessment of the strengths and weaknesses of the technique is provided, and suggestions made for further improvements and research.     

Disturbance and species displacement: different tolerances to stream drying and desiccation in a native and an invasive crayfish

1. Crayfish are among the most threatened taxa in the world and invasive crayfish are the primary cause of the decline of native crayfish. Most research has emphasised biotic interactions as the mechanism by which native crayfish are displaced by invasives, although crayfish occupy variable environments and the role of disturbance in facilitating crayfish invasion and displacement is understudied. 2. We compared tolerance to a disturbance, stream drying, in a native and invasive crayfish as a potential mechanism to explain their distribution. Our experiments and observations were conducted across scales, from laboratory environmental chambers to stream mesocosms to field sampling. We hypothesised that the invasive crayfish would be more tolerant of desiccation than the native, and that this physiological distinction between the two would be reflected in their distribution in relation to stream drying. 3. In the laboratory, the native crayfish Orconectes eupunctus was less tolerant of desiccation than the invasive Orconectes neglectus chaenodactylus, with all native crayfish dying within 2 days without water, while some of the invasive crayfish survived for nearly 2 weeks. Under simulated stream drying in mesocosms, only the native O. eupunctus survived less well than in a control. Field sampling demonstrated a significant negative relationship between O. eupunctus density and low summer flows, while O. neglectus density was positively associated with low summer flows. The greater resistance of O. neglectus to drying could, through priority effects, inhibit recolonisation by O. eupunctus once flow resumes. 4. Abiotic disturbances are potentially important to the displacement of native by invasive crayfish. Disturbance mediated displacement of aquatic species provides both an opportunity to conserve native species by maintaining or restoring habitat and disturbance regimes and is also a challenge due to increasing human water demand, flow regime alteration and global climate change.     

Effects of pulsed and pressed disturbances on the benthic invertebrate community following a coal spill in a small stream in northeastern USA

In September 1999, a coal-carrying train derailed and spilled 180,000–270,000 kg of coal into the Cayuga Inlet near Ithaca, New York. This study determined the immediate effects of the coal spill and the clean up procedures on the aquatic invertebrate community, and whether the stream recovered from this event after 2 years. Benthic invertebrate samples were taken both upstream and downstream of the coal spill immediately following clean-up efforts and two years later. Just after the coal spill, the total abundance and species richness of aquatic invertebrates were significantly lower downstream of the spill, suggesting that the disturbance caused increased mortality and/or emigration compared to a reference site upstream. Taxa affected most were grazers and turbidity-susceptible invertebrates. Two years later the invertebrate communities were similar upstream and downstream of the spill, except for an increase in the percent of the dominant genus, Hydropsyche(Trichoptera: Hydropsychidae). We speculate that long-term effects of channelization of the stream that occurred during the clean-up prevented the invertebrate assemblage from returning to the conditions observed in a reference site upstream of the coal spill. We propose that large scale environmental clean-ups should be designed to avoid altering ecosystems permanently, and that streams should be allowed to recover naturally without destructive human intervention.     

Effects of floods versus low flows on invertebrates in a New Zealand gravel-bed river

1. Floods and low flows are hydrological events that influence river ecosystems, but few studies have compared their relative importance in structuring invertebrate communities. Invertebrates were sampled in riffles and runs at eight sites along 40 km of a New Zealand gravel‐bed river every 1–3 months over 2.5 years, during which time a number of large flood and low flow events occurred. Flows were high in winter and spring, and low in summer and autumn. Four flow‐related variables were calculated from hydrological data: flow on the day of sampling (Q_sample), maximum and minimum flow between successive samples (Q_max and Q_min, respectively), and the number of days since the last bed‐moving flood (N_days). 2. The invertebrate community was summarised by relative densities of the 19 most abundant taxa and four biotic metrics [total abundance, taxon richness, the number of Ephemeroptera, Plecoptera and Trichoptera taxa (i.e. EPT richness), and per cent EPT]. Invertebrate density fluctuated greatly, and was high in summer and autumn, and low during winter and spring. Stepwise multiple regression (SMR) analysis was used to investigate relationships between the invertebrate community and season, flow, habitat and water temperature. 3. Seasonal variables were included in almost 50% of the SMR models, while flow‐related variables were included in >75% of models. Densities of many taxa were negatively correlated to Q_min and Q_max, and positively correlated to N_days, suggesting that while high flows reduced invertebrate densities, densities recovered with increasing time following a flood. Although season and flow were confounded in this study, many of the taxa analysed display little seasonal variation in abundance, suggesting that flow‐related variables were more important in structuring communities than seasonal changes in density associated with life‐cycles. 4. Five discrete flood and low flow events were identified and changes to invertebrate communities before and after these events examined. Invertebrate densities decreased more commonly after floods than after low flows, and there was a significant positive relationship between the number of taxa showing reductions in density and flood magnitude. Densities of most invertebrates either remained unchanged, or increased after low flow events, except for four taxa whose densities declined after a very long period (up to 9 months) of low flow. This decline was attributed to autogenic sloughing of thick periphyton communities and subsequent loss of habitat for these taxa. 5. Invertebrate communities changed more after floods and the degree of change was proportional to flood magnitude. Community similarity increased with increasing time since the last disturbance, suggesting that the longer stable flows lasted, the less the community changed. These results suggest that invertebrate communities in the Waipara River were controlled by both floods and low flows, but that the relative effects of floods were greater than even extended periods of extreme low flow. 6. Hydraulic conditions in riffles and runs were measured throughout the study. Riffles had consistently faster velocities, but were shallower and narrower than runs at all measured flows. Invertebrate density in riffles was expressed as a percentage of total density and regressed against the flow‐related variables to see whether invertebrate locations changed according to flow. Significant negative relationships were observed between the per cent density of common taxa in riffles and Q_sample, Q_max and Q_min. This result suggests either that these animals actively drifted into areas of faster velocity during low flows, or that their densities within riffles increased as the width of these habitats declined.     

Size‐related and diel variations in microhabitat use of three endangered small fishes in a Mediterranean coastal stream

This study analysed the microhabitat use of three endangered fish species, Andalusian toothcarp Aphanius baeticus , Iberian loach Cobitis paludica and sand smelt Atherina boyeri , in a coastal stream stretch. Plastic minnow traps were set both during daytime and at night on the bottom and at the surface. Depth and presence of effective refuge were recorded for each trap. To assess size‐related changes in microhabitat use individuals of each species were classified in three size classes. The three species preferentially used bottom positions in the water column, though this behaviour was more evident in the case of Iberian loach. While large Iberian loach remained active at night Andalusian toothcarp and sand smelt were strongly diurnal, especially larger individuals. The three species showed a clear ontogenetic change in microhabitat preferences towards deeper waters. Small Andalusian toothcarp and medium‐sized Iberian loach used deeper microhabitat in the presence of refuge. Large Andalusian toothcarp consistently preferred exposed microhabitat at any time. Andalusian toothcarp using refuge were smaller at any time, while Iberian loach followed this pattern only at night. The differential vulnerability of these species to different predators (aerial and aquatic; diurnal and nocturnal) could explain the observed patterns in microhabitat use. Fish tended to co‐occur in microhabitats either due to habitat characteristics independently of species or due to species independently of habitat. Andalusian toothcarp segregated spatio‐temporally from sand smelt and Iberian loach, but these species occurred independently of each other. According to these co‐occurrence patterns, Andalusian toothcarp would be more sensitive than Iberian loach or sand smelt to interspecific interactions.     

Time spent in the drift by downstream-dispersing invertebrates in a Lake District stream

1. The objective was to determine the time spent in the drift by different taxa of stream invertebrates. Most data were obtained from an earlier experimental study to determine the distances travelled by drifting invertebrates of 16 taxa in Wilfin Beck. Experiments were performed at two sites: ‘site 4’ in a stony, fast‐flowing, section of stream, ‘site 3’ in a deeper stream section where macrophytes were abundant. 2. The significant relationship between the mean distance x (m) travelled in the drift and modal water velocity V (m s^?1) was described by a power function in the earlier study but, as the power was close to one, a linear relationship has now been found to provide a satisfactory model. The rate of increase in x (m) with increasing V varied considerably between taxa. The mean time [mean t (s)] spent in the drift was estimated by dividing each x (m) by the appropriate V. Mean t (s) for each taxon was usually very constant over a wide range of V at each site (0.10–0.60 m s^?1 at site 4, 0.15–0.53 m s^?1 at site 3). A simple model estimated the time spent in the drift by different percentages (e.g. 75, 50, 10 and 1%) of the drifting invertebrates. 3. The experimental taxa at site 4 were divided into three groups according to the mean time spent in the drift. Mean t (s) for the five taxa in group 1 (32.8 s) was not significantly different from that obtained in control experiments with a mixed group of dead invertebrates. A similar time (33.0 s) was obtained for the five taxa in group 2, except at water velocities less than 0.2 m s^?1 when the mean t (s) decreased to 15–21 s. Mean t (s) was constant for each of the six taxa in group 3, and significantly less than that for groups 1 and 2. Mean values ranged from 28.8 s for Ephemerella ignita to only 9.4 s for Baetis rhodani and Gammarus pulex. All mean values were lower at site 3, presumably because of the dense stands of macrophytes, with mean values of 12.9 s for the five taxa in group 1 (equalling the value for dead invertebrates). Mean values for the six remaining taxa varied from 6.4 s for Simulium spp. to only 4.9 s for Baetis rhodani and 4.8 s for Gammarus pulex. It was concluded from a discussion of this study that the time spent in the drift may provide a useful measure for comparing the downstream dispersal of invertebrates in different streams, and may be a useful addition to models for the drift feeding of salmonids.     

Habitat heterogeneity drives the geographical distribution of beta diversity: the case of New Zealand stream invertebrates

To define whether the beta diversity of stream invertebrate communities in New Zealand exhibits geographical variation unexplained by variation in gamma diversity and, if so, what mechanisms (productivity, habitat heterogeneity, dispersal limitation, disturbance) best explain the observed broad‐scale beta diversity patterns. We sampled 120 streams across eight regions (stream catchments), spanning a north–south gradient of 12° of latitude, and calculated beta diversity (with both species richness and abundance data) for each region. We explored through a null model if beta diversity deviates from the expectation of stochastic assembly processes and whether the magnitude of the deviation varies geographically. We then performed multimodel inference analysis on the key environmental drivers of beta diversity, using Akaike's information criterion and model and predictor weights to select the best model(s) explaining beta diversity. Beta diversity was, unexpectedly, highest in the South Island. The null model analysis revealed that beta diversity was greater than expected by chance in all eight regions, but the magnitude of beta deviation was higher in the South Island, suggesting differences in environmental filtering and/or dispersal limitation between North and South Island. Habitat heterogeneity was the predominant driver of beta diversity of stream macroinvertebrates, with productivity having a secondary, and negative, contribution. This is one of the first studies accounting for stochastic effects while examining the ecological drivers of beta diversity. Our results suggest that local environmental heterogeneity may be the strongest determinant of beta diversity of stream invertebrates, more so than regional‐ or landscape‐scale variables.     

Species composition,abundance and emergence phenology of midges (Deiptera: Chironomidae) in a brown-water stream of West-Central Alberta,Canada

Sixteen floating, box type emergence traps, each covering 0.1 m², were placed along 150 m of a third-order stretch of the Bigoray River, a slow-flowing, vegetation-choked, brown-water muskeg stream. Effects of trap design, trap shading, length of sampling interval, and stream velocity on the number of midges caught was examined. Of the 112 species of Chironomidae caught, 32 species made up 90% of the catch and their emergence phenology is described in detail. There were more rare species than expected from Preston's lognormal distribution. Percentage of Bigoray species belonging to Tanypodinae, Orthocladiinae, Chironomini and Tanytarsini was 18%, 43%, 20%, and 19%, and was compared with 27 other studies on lotic chironomids. During the 140-day emergence period an average of 19.3 × 10³ chironomids emerged per square meter of stream. Based on changes in male:female ratios throughout a species emergence period, it was postulated that 30% of the Bigoray chironomid species were univoltine, 50% were bivoltine, and 20% were trivoltine.     

Drought intensification alters the composition,body size,and trophic structure of invertebrate assemblages in a stream mesocosm experiment

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