The longitudinal distribution of riverine benthos: A river dis-continuum?

We used several techniques to analyze 10–11 collections from each of 13 stations along a mountain stream gradient to examine the hypothesis that there was a statistically significant (p < 0.05) uniform downstream gradient in benthic distribution. Random skewer techniques suggested that there was a significant downstream gradient of individuals per species, but only a weak gradient in species per station, and no gradient in functional guild composition. Ordination and clustering of either taxonomic or functional guild data suggested the existence of four longitudinally-defined groups of stations. Rarefaction curves also implied longitudinal groupings, with differences among groups implying species replacement than changes in organism numbers. Two species replacement indexes confirmed that suggestion. In summary, the distribution of the benthos in the creek may best be characterized as a punctuated gradient where gradual downstream changes in community composition are punctuated by sudden shifts to new community types. This punctuated gradient appears caused by changes in stream physical and chemical characteristics; the rapid nature of the changes hampers the interpretation of longitudinal analyses. Techniques such as random skewers and cluster analysis of rarefied samples will prove helpful in discerning pattern in stream benthos.     

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

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

Dynamics of Invertebrate Benthic Communities and Drift in a Regulated River of Central Spain

We document invertebrate benthic and drift dynamics in a regulated river in central Spain at two temporal scales: seasonal (for both benthos and drift) and daily (for drift). The benthic abundance of individuals and taxon richness generally increased in the summer. Drift abundance showed no seasonal or daily variation, but taxon richness of drifting individuals was higher in the spring. Both ben‐thos and drift showed clear seasonal changes in taxonomic composition. Interestingly, some benthic taxa showed their highest abundances in the spring, while others were more abundant in the summer. In contrast, most drifting taxa were more abundant in the spring. Different functional feeding groups showed different patterns of variation throughout the year, both in the benthos and the drift. Daily variations in drift were present in very few taxa and functional feeding groups, and only in some seasons. (© 2005 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

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.     

Estimating processing time of stream benthic samples

Multiple regression analysis was used to generate equations relating time required to sort and tabulate benthic invertebrates to size characteristics of benthic samples collected from a 5th-order river and a 3rd-order woodland stream. Number of invertebrates in a sample was the primary determinant of sample sorting time. Amount of detritus, occurrence of filamentous algae (Cladophora) and source stream contributed significant but minor additional variability to estimates (cumulativeR ² = 0.95). When number of animals was excluded as a variable, amount of detritus and presence/absence ofCladophora alone could be used to predict sorting time (R ² = 0.81). A typical sample containingCladophora required 29% longer to sort than samples containing equivalent amounts of organic material but noCladophora. The influence of sampler size and subsampling on processing and sorting time are considered. A general equation was derived to provide guidelines for selecting a size of stream sampler that subsequently minimizes total processing time required to estimate density of benthos with acceptable, constant precision. Overall sample processing and sorting time are reduced by using a smaller sampler or by subsampling only if benthic densities of animals are high. Use of regression equations to anticipate processing and sorting time required for a particular sampling program permits development of more efficient designs.     

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.     

Feeding ecology of a vertebrate assemblage inhabiting a stream of NW Spain (Riobo; Ulla basin)

The food resource use of a stream in NW Spain by fish (Salmo trutta L. and Anguilla anguilla L.), birds (Cinclus cinclus L. and Motacilla cinerea L.) and mammals (Galemys pyrenaicus G. and Neomys anomalus C.) was studied. Data on seasonal diets and stream benthos prey were used to determine prey selection patterns.Caddisfly larvae are the main resource for Cinclus and Galemys, but these predators also consumed other benthic prey. Salmo fed on a wide range of benthic invertebrates, emergent pupae and terrestrial prey, whereas Anguilla consumed primarily benthic invertebrates, especially Lumbricids. Neomys fed mainly on terrestrial prey (Gasteropods and Lumbricids), but also consumed aquatic prey. Motacilla captured aquatic insects both in larval and aerial stages, as well as terrestrial prey.Both prey availability and selection led to seasonal differences in the use of food resources. All species showed a marked prey selection of aquatic taxa. Prey size plays an important role in this selection, most species consuming the largest of available prey sizes. In spite of the fact that all species feed upon freshwater invertebrates, substantial resource partitioning was observed in all seasons. This partitioning may be attributable to morpholological and physiological differences. Nevertheless, Anguilla and Galemys, two quite different animals, did feed on the same prey much of the time.     

Concordance among stream assemblages and spatial autocorrelation along a fragmented gradient

Patterns of spatial autocorrelation of biota and distributional similarity (concordance) between assemblages of different organism groups have important implications in both theoretical ecology and biodiversity conservation. Here we report environmental gradients and spatial distribution patterns of taxonomic composition among stream fish, benthic macroinvertebrate, and diatom assemblages along a fragmented stream in south‐western France. We quantified spatial patterns of lotic assemblage structure along this stream, and we tested for concordance in distribution patterns among the three taxonomic groups. Our results showed that both environmental characteristics and stream assemblages were spatially autocorrelated. For stream fish and diatom assemblages, these patterns reflected assemblage changes along the longitudinal stream gradient, whereas environmental variables and benthic macroinvertebrates exhibited a more patchy spatial pattern. Cross‐taxa concordance was significant between stream fish and diatoms, and between stream fish and benthic macroinvertebrates. The assemblage concordance between stream fish and diatoms could be attributed to similar responses along the longitudinal gradient, whereas those between stream fish and benthic macroinvertebrates may result from biotic interactions. Based on potential dispersal capacities of taxa, our results validated the hypotheses that weakly dispersing taxa exhibit greater concordance than highly dispersing ones and that dispersal capacities affect how taxonomic groups respond to their local environment. Both diatoms and highly dispersing stream fish were affected by stream fragmentation (i.e. the number of dams between sites), while the effect of fragmentation was not significant for invertebrates that fly well in their adult stage, thus emphasizing the importance of the way of dispersal. These results suggest that addressing the effects of dispersal capacity on stream assemblage patterns is crucial to identifying mechanisms behind patterns and to better understanding the determinants of stream biodiversity.     

Response of Benthic Invertebrates to Disturbance From Stream Restoration: The Importance of Bryophytes

The response of benthic invertebrates to disturbance from stream restoration was studied in the Reinikankoski rapids, central Finland. I hypothesized that stream bryophytes could act as a refugium for invertebrates. The restoration procedure destroyed nearly half of the bryophytes present at the study reach, and invertebrate densities decreased sharply immediately after the restoration. Within 2 weeks, invertebrates had recolonized the disturbed reach, and within 1 month peak numbers were attained. Invertebrates showed a clear association with bryophytes, especially after the restoration. My study shows that invertebrate recovery can be relatively fast in winter, and underlines the importance of stream mosses as invertebrate habitat and refugia. They should be taken in consideration in stream restoration projects by leaving patches of stream bottom intact.     

Riffle-pool geomorphology disrupts longitudinal patterns of stream benthos

An Ozark Plateau stream was studied to determine the influence of distinct pool and riffle geomorphology on the longitudinal zonation of macroinvertebrate species assemblages and functional group classification. All study sites were dominated by alluvial pool and riffle channel form and the first two orders became intermittent during summer months. Nine benthos samples were collected seasonally from riffles and pools at each of five sites using a vacuum benthos sampler. Diel temperature pulse and coarse particulate organic matter (CPOM) were measured at each site also. Water temperature was most variable in second order, and CPOM on riffles was not more abundant in upstream reaches. Annual average density and biomass of invertebrates were highest in third and fourth orders, respectively. Diversity was depressed in the intermittent headwaters sites. Macroinvertebrate functional groups did not exhibit strong longitudinal trends as predicted by the river continuum model, with species assemblages apparently more strongly affected by the segment-level physical template, although shredders were more abundant in the headwaters during fall and winter. This study indicates that a reach-level perspective based on channel form is a necessary complement to holistic stream ecosystem models, especially in alluvial gravel streams.     

An ecological investigation of the Portneuf River, Idaho: a semiarid-land stream subjected to pollution

The Portneuf River (mean monthly discharge near mouth 1.4-15.4 m³/s) in southeastern Idaho, U.S.A., flows through an area of reduced precipitation (approx. 33 cm/year) and is dependent on underground aquifers and snowmelt runoff from the surrounding mountains for its water. The stream was examined at ten locations, distributed over its 156-km course, during the period 1967-1971. The Portneuf River is shown to have undergone a number of changes from its natural state as evidenced by alterations in water quality and the distribution of benthic invertebrates along the stream course. Of particular interest are changes brought about by the use of the stream for irrigation and by runoff from agricultural lands, factors whose effects are magnified by the semiarid conditions of the region and by poor soil-conservation practices. However, the stream also is affected by wastes from a sewage-treatment plant, phosphate-processing operations, and an assortment of scattered urban sources. Benthic invertebrates were collected during all four seasons by means of artificial substratum samplers and during summer and autumn by a qualitative dip-net technique. In general, the samplers were more effective in obtaining a representative picture ofthe fauna. However, neither procedure alone gave as much information as the combined results. The artificial substratum collections are not believed to be representative of the usual effects of stream devvatering by irrigation withdrawal in as much as the samplers provide refugia for the benthos during the periods of reduced habitat.     

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.     

Recent land‐use changes affect stream ecosystem processes in a subtropical island in Brazil

Land‐use changes such as conversion of natural forest to rural and urban areas have been considered as main drivers of ecosystem functions decline, and a large variety of indicators has been used to investigate these effects. Here, we used a replicated litter‐bag experiment to investigate the effects of land‐use changes on the leaf‐litter breakdown process and leaf‐associated invertebrates along the forest–pasture–urban gradient located in a subtropical island (Florianópolis, SC, Brazil). We identified the invertebrates and measured the litter breakdown rates using the litter bags approach. Litter bags containing 3 g of dry leaf of Alchornea triplinervia were deployed on forest rural and urban streams. Principal component analysis, based on physico‐chemical variables which, confirmed a gradient of degradation from forest to urban streams with intermediate values in rural areas. In accordance, shredder richness and abundance were lower in rural and urban than in forest streams. The land‐use changes led also to the dominance of tolerant generalist taxa (Chironomidae and Oligochaeta) reducing the taxonomic and functional diversity in these sites. Leaf‐litter breakdown rates decreased from forest to rural and finally to urban areas and were associated with changes in pH, water velocity, dissolved oxygen and abundance of leaf‐shredding invertebrates, although global decomposition rates did not differ between rural and urban streams. Overall, this study showed that land‐use changes, namely to rural and urban areas, have a strong impact on tropical streams ecosystems, in both processes and communities composition and structure. Despite of being apparently a smaller transformation of landscape, rural land use is comparable to urbanisation in terms of impact in stream functioning. It is thus critical to carefully plan urban development and maintain forest areas in the island of Florianópolis in order to preserve its natural biodiversity and aquatic ecosystems functioning.     

Macrozoobenthos of the Small Aral Sea

Benthic assemblages of the Small Aral Sea were studied in 1999. The species composition, density, and biomass of benthic invertebrates were determined. The vertical distribution pattern and its dynamics in the course of desiccation of the sea were analyzed. It is shown that the vertical distribution of the benthos has changed substantially over the last 30 years. The biomass of the benthos in the inshore zone increased, while in the central part of the sea, benthic invertebrate populations declined and gradually disappeared. These changes are likely to be due to the accumulation of organic matter in the bottom sediments and the weakening of hydrodynamic processes, which could lead to oxygen depletion in deep water. In the 1990s, the benthic assemblages of the Small Aral Sea were relatively stable and highly productive. It is concluded that the Aral Sea is a promising area for fishery restoration.     

Using Diatom Assemblages to Assess Urban Stream Conditions

We characterized changes in diatom assemblages along an urban-to-rural gradient to assess impacts of urbanization on stream conditions. Diatoms, water chemistry, and physical variables of riffles at 19 urban and 28 rural stream sites were sampled and assessed during the summer base flow period. Near stream land use was characterized using GIS. In addition, one urban and one rural site were sampled monthly throughout a year to assess temporal variation of diatom assemblages between the urban and rural stream sites. Canonical correspondence analysis (CCA) showed that the 1st ordination axis distinctly separated rural and urban sites. This axis was correlated with conductivity (r = 0.75) and % near-stream commercial/industrial land use (r = 0.55). TWINSPAN classified all sites into four groups based on diatom assemblages. These diatom-based site groups were significantly different in water chemistry (e.g., conductivity, dissolved nutrients), physical habitat (e.g., % stream substrate as fines), and near-stream land use. CCA on the temporal diatom data set showed that diatom assemblages had high seasonal variation along the 2nd axis in both urban and rural sites, however, rural and urban sites were well separated along the 1st ordination axis. Our results suggest that changes in diatom assemblages respond to urban impacts on stream conditions.     

Are algal communities driven toward maximum biomass?

In this continental-scale study, we show that in major benthic and planktonic stream habitats, algal biovolume--a proxy measure of biomass--is a unimodal function of species richness (SR). The biovolume peak is observed at intermediate to high SR in the benthos but at low richness in the phytoplankton. The unimodal nature of the biomass-diversity relationship implies that a decline in algal biomass with potential harmful effects on all higher trophic levels, from invertebrates to fish, can result from either excessive species gain or species loss, both being common consequences of human-induced habitat alterations. SR frequency distributions indicate that the most frequent richness is habitat-specific and significantly higher in the benthos than in the plankton. In all studied stream environments, the most frequent SR is lower than the SR that yields the highest biovolume, probably as a result of anthropogenic influences, but always within one standard deviation from it, i.e. they are statistically indistinguishable. This suggests that algal communities may be driven toward maximum biomass.     

Upstream-downstream movements of aquatic invertebrates in a Rocky Mountain stream

Simultaneous collections of drift and organisms moving either upstream or downstream in association with the substrate were made using a specially designed sampler. Samples were taken in a diel series along a transect across the study riffle of a Colorado foothills stream on six dates over an annual cycle. In addition to longitudinal movements, taxonomic composition and diel periodicity were evaluated. The insect-dominated fauna showed a net downstream displacement. Only the caddisflies Helicopsyche borealis and Hesperophylax occidentalis exhibited net upstream movement, primarily a result of low drift frequencies. The taxonomic composition of moving invertebrates differed from that of the benthos. Drift resembled downstream moving substrate-associated invertebrates in composition, but differed from that of the upstream directed fauna. Taxa collectively exhibited four types of diel patterns: 1) similar downstream (drift and substrate-associated movements) patterns, which generally differed from the upstream pattern; 2) similar benthic (upstream and downstream) patterns, which differed from that of drift; 3) aperiodic patterns; and 4) independent patterns for each type of directional movement. Analysis of size classes based on head capsule width for the mayfly Baetis tricaudatus showed significantly smaller size in stationary individuals compared with moving individuals in the population and revealed that nymphs moving during the day were smaller than those moving at night.     

Vertical distribution of benthic community responses to fish predators, and effects on algae and suspended material

The vertical positioning of benthic invertebrates should be a trade-off between the risky, but productive, sediment surface and the safer, but physiologically harsher, conditions deeper down in the sediment. This is because the foraging efficiency of benthic fish decreases with sediment depth, whereas the sediment surface is generally better oxygenated and has a higher resource quality than lower layers. We studied how two benthic fish predators, bream (Abramis brama) and ruffe (Gymnocephalus cernuus), affected the community composition and vertical distribution of benthos, and their indirect effects on algae and suspended material, in field enclosures. Whereas bream had significant effects on the density, composition and distribution of the benthos, ruffe had no such effects. The total benthos biomass in bream treatments was an-order of magnitude lower in the upper sediment layer (0–1 cm) and three times lower in the middle layer (1–3 cm) than in the controls, whereas there were no significant effects in the deepest layer (3–10 cm). Bivalves persisted in the deepest layer although their density was reduced in shallow sediment, whereas gastropods faced the risk of local extinction in the presence of bream. As indirect effects, small-bodied cladocerans, phytoplankton, periphyton and both organic and inorganic suspended material were higher in the bream treatments. We␣conclude that the impact of bream diminished substantially with increasing sediment depth, enabling invertebrates to survive in the sediment and to persist in the presence of bream. However, there were␣no␣indications of any group adjusting their vertical position behaviourally as a response to predation threat.