Habitat structure,resources and diversity: the separate effects of surface roughness and macroalgae on stream invertebrates

Habitat structure has pervasive effects on community composition and diversity, with physically complex habitats often containing more species than physically simple ones. What factors or mechanism drive this pattern is little understood, but a complicating problem is that different sources of habitat structure can be confounded in both surveys and experiments. In this study, we carried out an experiment in which two sources of habitat structure, attached macroalgae and substrate surface texture, were separately manipulated to discern their joint and separate effects upon the diversity and composition of colonizing macroinvertebrates in a stony, upland stream. Because stream algae vary markedly in abundance in both space and time, we also sampled the epilithon of stream stones at two spatial scales on eight dates over 2 years to gain some preliminary data on how stream algae vary between individual substrata over time. Experimental substrata had either a smooth (siltstones, sandstones, crystal-poor felsic volcanics, plain paving bricks) or rough (granodiorites, crystal-rich felsic volcanics, sand-blasted paving bricks) surface. We allowed these substrata to be colonized naturally by macroalgae, mostly the filamentous red alga Audouinella hermannii. Half of each of the rough and smooth substrata were selected at random and the macroalgae gently sheared off. All substrata were defaunated with a household insecticide with little field persistence, set out randomly through the study riffle, and invertebrates allowed to colonize them for 14 days. Some substrata were sampled immediately to check the efficacy of faunal and algal removals, which proved to be successful. Experimental results showed that both surface texture and macroalgae increase species richness independently of each other. Surface texture had no effect on densities, while macroalgae increased colonization densities, but rarefaction showed that both sources of habitat structure increased species richness above values expected simply on the basis of the numbers of colonists. However, reference stones with high macroalgal cover had the same species richness as those with low cover, suggesting that the effects of macroalgae on species richness are transient relative to those associated with surface texture. Epilithon samples taken at different times suggest that the magnitude of spatial variation in plant growth alters with time. If plants generally recolonize rough surfaces more quickly than smooth, then the effects of habitat structure on macroinvertebrates ought to be strongest after major disturbances during growing seasons of plants. Received: 1 September 1999 / Accepted: 10 January 2000     

You don’t belong here: explaining the excess of rare species in terms of habitat,space and time

Ecological communities are composed of a few common and several rare species. Many studies have evaluated the shape of abundance distribution curves, but few studies have assessed the causes of rarity. Using a dataset of stream macroinvertebrates, we investigated whether the excess of rare species in three focal communities of stones in riffles were common 1) in other habitats at the same stream site and period of sampling (environment), 2) in other stream sites in the same habitat and period of sampling (space), and 3) in other years in the same stream site and habitat (time). We observed that around 28% of the rare species were common in other habitats (environment), stream sites (space) or years (time). Among the three factors, rarity was mostly explained by habitat type, whereas a significant portion of the rare species in riffles were common in pools, submerged roots of terrestrial plants or in partially submerged moss patches. This result suggests that the presence in non‐optimum habitat is a strong determinant of the rarity observed in natural communities and most rare species are due to sampling artifacts or accidentally sampled transient species.     

Hierarchical structure of stream ecosystems: consequences for bioassessment

It has long been recognized that communities and their ecosystems are structured at several, nested spatial scales. But identifying the appropriate scale(s) to collect, analyse and interpret data to answer specific questions about ecosystems has been a vexing problem for ecologists. We collected observations of the benthic invertebrate community and its environment in 10 primarily agricultural tributary streams of the Thames River in southwestern Ontario, Canada. Within each stream we sampled two reaches, in each reach we sampled three riffles, and in each riffle we took three kick samples of invertebrates and characterized the substrate environment. We also characterized the habitat at each of the 20 reaches (10 streams × 2 reaches/stream). Most of the variability in the stream invertebrate community structure (as described with taxonomic richness and the biotic index of tolerance, as well as by the Bray-Curtis distance of the community composition from the mean at a spatial scale) was at larger spatial scales of among streams and between riffles. Much of the substrate and habitat variation was also at the larger spatial scales, as were correlations between the biota and the environment of the benthic invertebrate community. We concluded that for the purposes of bioassessment, characterization of one reach per stream is sufficient, at least in this context, for describing a stream and evaluating its health. Handling editor: R. Norris     

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

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

Spatial scale and the diversity of macroinvertebrates in a Neotropical catchment

1. Lotic ecosystems can be studied on several spatial scales, and usually show high heterogeneity at all of them in terms of biological and environmental characteristics. Understanding and predicting the taxonomic composition of biological communities is challenging and compounded by the problem of scale. Additive diversity partitioning is a tool that can show the diversity that occurs at different scales. 2. We evaluated the spatial distribution of benthic macroinvertebrates in a tropical headwater catchment (S.E. Brazil) during the dry season and compared alpha and beta diversities at the scales of stream segments, reaches, riffles and microhabitats (substratum types: gravels, stones and leaf litter). We used family richness as our estimate of diversity. Sampling was hierarchical, and included three stream segments, two stream reaches per segment, three riffles per reach, three microhabitats per riffle and three Surber sample units per microhabitat. 3. Classification analysis of the 53 families found revealed groups formed in terms of stream segment and microhabitat, but not in terms of stream reaches and riffles. Separate partition analyses for each microhabitat showed that litter supported lower alpha diversity (28%) than did stones (36%) or gravel (42%). In all cases, alpha diversity at the microhabitat scale was lower than expected under a null model that assumed no aggregation of the fauna. 4. Beta diversity among patches of the microhabitats in riffles depended on substratum type. It was lower than expected in litter, similar in stone and higher in gravel. Beta diversities among riffles and among reaches were as expected under the null model. On the other hand, beta diversity observed was higher than expected at the scale of stream segments for all microhabitat types. 5. We conclude that efficient diversity inventories should concentrate sampling in different microhabitats and stream sites. In the present study, sampling restricted to stream segments and substratum types (i.e. excluding riffles and stream reaches) would produce around 75% of all observed families using 17% of the sampling effort employed. This finding indicates that intensive sampling (many riffles and reaches) in few stream segments does not result in efficient assessment of diversity in a region.     

The influences of habitat and seasonal sampling regimes on the ordination and classification of macroinvertebrate assemblages in the catchment of the River Wye,Wales

Assemblages of benthic macroinvertebrates were assessed in relation to environmental variables at 45 sites in the catchment of the Welsh River Wye in 1982.Samples were collected from three habitats (margins, riffles, flats) and in two seasons (Spring and Autumn), separated and combined data sets being ordinated by DECORANA and classified by TWINSPAN. Correlations between primary ordinations were high (p < 0.001) and always strongly related to total hardness or pH; secondary ordinations were also intercorrelated and were related to stream slope.In each instance, total hardness and slope were the strongest discriminants between site groupings produced by TWINSPAN. It is concluded that relationships between faunal assemblages, stream chemistry and stream physiography would be effectively detected in the Wye catchment in either Spring or Autumn and in each of the habitats sampled. However, the most precise categorization of assemblage type required a sampling strategy which combined habitat and seasonal data.     

The ecology of two intermittent streams in Victoria, Australia

1. We compared aquatic macroinvertebrate assemblage composition within and between habitats (pools and riffles) at four sites on two intermittent streams in Victoria, Australia during a drought year followed by a wetter year to see how different spatial and temporal scales influenced patterns of community structure detected by multivariate techniques of ordination and classification. We also examined the fit between faunal groups and ‘flow phases’ characterized by multivariate analyses of physicochemical variables in an earlier paper. 2. Pools contained more taxa and individuals than riffles. At the more temporary site, there was considerable overlap between the faunal composition of pools and riffles, whereas habitat-specific assemblages consistently occurred at the more permanent sites. Most taxa were either tolerant, permanent stream forms able to persist in streams that dry briefly, or facultative species that occur in lotic or lentic habitats. Few species appeared specifically adapted to temporary waters. 3. Within habitats, there were some matches between faunal groups and flow phases, especially with the onset of the ‘diminishing flow’ phase, characterized by rising water temperature and conductivity, and declining pH and dissolved oxygen. However, disjunctions among faunal groups were less marked than those between flow phases, and were absent at an intermediate scale when habitats were pooled within sites. At the broadest scale of analysis (within habitats between sites and years), historical events (e.g. whether pools upstream dried completely during the previous summer) apparently influenced community composition as much as site-specific abiotic differences.     

Spatial scale impacts microbial community composition and distribution within and across stream ecosystems in North and Central America

A mechanistic understanding of factors that structure spatiotemporal community composition is a major challenge in microbial ecology. Our study of microbial communities in the headwaters of three freshwater stream networks showed significant community changes at the small spatial scale of benthic habitats when compared to changes at mid- and large-spatial scales associated with stream order and catchment. Catchment (which included temperate and tropical catchments) had the strongest influence on community composition followed by habitat type (epipsammon or epilithon) and stream orders. Alpha diversity of benthic microbiomes resulted from interactions between catchment, habitat, and canopy. Epilithon contained relatively more Cyanobacteria and algae while Acidobacteria and Actinobacteria proportions were higher in epipsammic habitats. Turnover from replacement created ~60%–95% of beta diversity differences among habitats, stream orders, and catchments. Turnover within a habitat type generally decreased downstream indicating longitudinal linkages in stream networks while between habitat turnover also shaped benthic microbial community assembly. Our study suggests that factors influencing microbial community composition shift in dominance across spatial scales, with habitat dominating locally and catchment dominating globally.     

Littoral benthos of a Victorian lake and its outlet stream: Spatial and temporal variation

Abstract There have been few comparative studies of the fauna in the two major types of freshwater systems, lakes and streams, in the one locality. This study compared the faunal assemblages at two times of the year (summer and winter) on stones in three locations: the littoral zone of two shores in Lake Purrumbete (one wave-exposed and one sheltered) and riffles in the Curdies River, which flows out of the lake. The lake fauna was dominated by crustaceans, gastropods and planarians, whereas the stream fauna was dominated by insects. The most abundant lake taxa were also present, but much less abundant, in the stream. The total number of species and individuals and densities of some common species varied between the three locations and between seasons. However, no consistent pattern reflecting a difference between the two lake shores was evident. Interpretation of MDS ordinations of the lake fauna was dependent on the data standardizations applied, with increased separation of the season-location combinations when species were standardized to equal total abundances; the seasonal difference was always greatest for the exposed (cliff) shore. In contrast, MDS on the stream fauna showed seasonal differences under all standardizations. This was consistent with the high seasonal turnover of species in the stream compared with the lake. These results demonstrate that, even within a local area with similar geology and connected water bodies, lake and stream fauna from the same substratum (stones) can be markedly different. Taxa that occurred in both were more abundant in the lake, whereas seasonal differences in abundance were much greater in the stream.     

Scale and the detection of land-use effects on morphology, vegetation and macroinvertebrate communities of grassland streams

1. Land‐use studies are challenging because of the difficulty of finding catchments that can be used as replicates and because land‐use effects may be obscured by sources of variance acting over spatial scales smaller than the catchment. To determine the extent to which land‐use effects on stream ecosystems are scale dependent, we designed a whole‐catchment study of six matched pairs (pasture versus native tussock) of second‐order stream catchments, taking replicate samples from replicate bedforms (pools and riffles) in each stream. 2. Pasture streams had a smaller representation of endemic riparian plant species, particularly tussock grasses, higher bank erosion, a somewhat deeper layer of fine sediment, lower water velocities in riffles, less moss cover and higher macroinvertebrate biodiversity. At the bedform scale, suspendable inorganic sediment (SIS) was higher in pools than riffles and in pasture streams there was a negative relationship between SIS and the percentage of the bed free of overhanging vegetation. Differences between stream reaches (including any interactions between land use and stream pair) were significant for SIS, substrate depth and characteristics of riparian vegetation. There were also significant differences between replicate bedforms in the same stream reaches in percentage exotic species in overhanging vegetation, percentage moss cover, QMCI (Quantitative Macroinvertebrate Community Index – a macroinvertebrate‐based stream health index) and macroinvertebrate density. 3. Significant differences among stream reaches and among replicate bedform units within the same reach, as well as interactions between these spatial units and land‐use effects, are neither trivial nor ‘noise’ but represent real differences among spatial units that typically are unaccounted for in stream studies. Our multi‐scale study design, accompanied by an investigation of the explanatory power of different factors operating at different scales, provides an improved understanding of variability in nature.     

The spatial heterogeneity of diatoms in eight southeastern Ohio streams: how far does a single riffle reach?

Periphyton is a commonly used biomonitoring tool for streams. Often only one or few riffles are sampled and assumed to be representative of a stream reach. Current literature focuses on periphyton heterogeneity at small scales, on individual rocks within a riffle, and larger scales, within watersheds or ecoregions. The intermediate scales, within single riffles or among riffles, have not been adequately addressed. The purpose of this research was to determine how many riffles must be sampled in order to represent a reach and whether the number of necessary riffles varied with stream health. Since periphyton is sensitive to habitat change, it was hypothesized that heterogeneity would be primarily partitioned among riffles. Eight to ten consecutive riffles were sampled at eight individual stream reaches. Sampled reaches were categorized based on previously collected bioassesment data: three non-attaining, three partially-attaining, and two fully-attaining water quality standards as defined by the Ohio Environmental Protection Agency. Data were analyzed using the Bray-Curtis Similarity Index, Hill’s N₂ dominance diversity index, and the Acid Mine Drainage Diatom Index of Biotic Integrity. Diatoms appeared to be patchily distributed within a reach. This patchiness often led to varied relative abundance of common species and the introduction or loss of rare species among riffles. To account for this variation within a reach, at least two riffles should be sampled. However, a multimetric index may correctly classify a stream based on a one-riffle sample. Variation does not appear to correspond directly to stream health, but to species richness and diversity.     

Coarse Particulate Organic Matter Distribution in the Pools and Riffles of a Second-order Stream

Coarse particulate organic matter distribution was investigated in a 270-m interval of a 2nd-order forest stream, the Yanase River. The dominant tree type was Keyaki (Zelkova serrata). CPOM sampling was conducted, and the sampled CPOM was sorted into leaves and branches, and water depth and current velocity were measured along with stone width, stone height and distance between the stones in the riffles. The collected CPOM was categorized by their accumulation type: LSS, SLP, SLPi, SLPo and DD. LSS was leaf packs at the leading edge of the stones, SLP was the sinking leaf packs in the pools, SLPi was the sinking leaf packs at the inner side of the stream bend in the pools, while SLPo was the sinking leaf packs at the outer side of the stream bend, and finally, DD was the leaf packs in the debris dams. The accumulated CPOM amounts at each leaf pack type were correlated with the measured physical stream variables. From the results, CPOM accumulation in riffles is controlled by stones projecting above the water surface and by their arrangements. In pools, CPOM accumulation occurs at high stream flow where the pools provide low velocities and a thick boundary layer of relatively quiescent flow. At stream meandering points, CPOM accumulation occurs by production of a secondary flow. Finally, CPOM accumulation in debris dams is important where they occur frequently. LSS was the largest in CPOM amount, and the stones in the riffles were the most retentive structures. On the other hand, SLP and SLPo were the least CPOM amounts, and the pools in the mainstream and at the outer side of the stream bend were the least retentive stream morphologies.     

A Large-scale Comparative Analysis of Riffle and Pool Fish Communities in an Upland Stream System

Fishes were sampled in riffle and pool habitats at 74 upland localities in the Little River system, southeastern Oklahoma and southwestern Arkansas, U.S.A. I asked how these two habitat-defined communities differed with regard to species abundance and incidence patterns, and how these differences varied along othree environmental gradients: elevation, stream gradient, and stream size. Riffle and pool communities showed distinct and significant differences when ordinated in multivariate space defined by species abundance patterns. Sites with similar pool communities did not have similar riffle communities, and riffle and pool communities responded to environmental gradients in different ways. Elevation was the best predictor of pool community structure, whereas stream size was the best predictor of riffle communities. Overall, riffle habitats had fewer species than pool habitats and formed significant subsets of pool communities at 12 of 74 sites. I predicted that at small stream localities where riffles were unstable, riffle species would form subsets of the pool species communities, and both community types should show high similarities. The presence of faunal subsets was not associated with stream size, but faunal similarities were significantly higher at small stream localities. At the species level, 14 species were significantly associated with pool habitats, while only two were associated with riffle habitats. Riffle and pool communities, although linked by a continuous habitat gradient at the local scale, responded differently to large-scale environmental gradients. Local differences between these communities were predictable based on stream size.     

The Effect of Riffle-Scale Environmental Variability on Macroinvertebrate Assemblages in a Tropical Stream

In a tropical stream (at the Soberaní;a National Park, Panama), different environmental factors were quantified in riffle habitats (water characteristics: velocity, depth, turbulence, and direction; stone characteristics: surface area, sphericity, and degree of burial; and others: substrate type, and canopy cover). Characteristics of macroinvertebrate assemblages (mean density of individuals, mean taxon richness, and cumulative taxon richness in three stones at each riffle) were related to both mean values and variability of these environmental factors at riffle scale. Macroinvertebrate density was higher in shallow, fast flowing, stony riffles, with low variability in dominant substrate type. Taxon richness was also higher in shallow riffles with loose, not buried stones, and water direction more or less parallel to the bank. Environmental variability resulted as important as mean values of environmental factors to explain variation in macroinvertebrate assemblages. This is the first study, to our knowledge, that quantifies substratum variability and demonstrates its influence on macroinvertebrate assemblages in a tropical stream.     

Space–time interactions and invertebrate assemblage change in stream networks

Streams form hierarchical, dendritic physical networks, but relatively little is known about how this spatial structure affects community assembly. We investigated interactions between changes over time in macroinvertebrate assemblages and their distribution in space (the space–time interaction) in stream networks. Assemblages were sampled from every tributary, and every reach between tributaries, to determine effects of network position on assemblage composition, in four West Coast, South Island, New Zealand, headwater networks. Using canonical redundancy analysis, we found that macroinvertebrate assemblages were significantly spatially structured and species assemblages changed significantly between two sampling periods. The most important environmental variables (averaged over all AIC models) explaining change in assemblage composition were related to disturbance, local habitat/resources and habitat size. The lack of a significant interaction between space and time, however, indicated the spatial pattern of assemblages remained the same over time, regardless of changes in assemblage composition. Consistent spatial structuring could be the result of unchanging processes such as those arising from the universal nature of stream topology and hydrology acting both on habitat‐ and dispersal‐ related community processes. Thus, we conclude that although community assemblages changed over time, the spatial arrangement of communities could potentially be predicted from stream network topology and hydrology.     

Habitat-specific estimates of competition in stream salmonids: A field test of the isodar model of habitat selection

Summary The population densities of sympatric Atlantic salmon,Salmo salar and brook charr,Salvelinus fontinalis, were measured in riffle and pool stream habitats to test whether non-linear isodars, a multispecific model of habitat selection based on ideal distribution assumptions, could (1) predict the distribution of densities between habitats and (2) reproduce the processes postulated to underlie spatial segregation and species interactions in previous laboratory and field studies. The model provided a good fit to observed density patterns and indicated that habitat suitability declined non-linearly with increased heterospecific competitor densities. Competitive effects in riffles appeared to be due to exploitative resource use, with salmon always emerging as the superior competitor. No evidence was found for interference competition in riffles. In contrast, interspecific competition in pools seemed to occur through exploitation and interference. The specific identity of the superior competitor in pools depended on the density of both species; pools provided the charr with refuge from competition with the salmon, presumably through the adoption by the charr of density-dependent behaviours, such as schooling and group foraging, that mitigated the negative impact of the salmon. Charr were displaced from the riffles toward the pools as the total salmon density increased. The isodar analysis, based on limited density data, successfully reproduced the processes suggested to underlie spatial segregation in previous field and laboratory studies and provided new insights into how changes in competitor densities modify habitat suitability in this system.     

Habitat overlap between predatory benthic fish and their invertebrate prey in streams: the relative influence of spatial and temporal factors on predation risk

1. The spatial heterogeneity of ecosystems as well as temporal activity patterns of organisms can have far‐reaching effects on predator–prey relationships. We hypothesised that spatiotemporal constraints in mesohabitat use by benthic fish predators would reduce habitat overlap with benthic invertebrates and lead to mesohabitat‐specific predation risks. 2. We analysed the spatiotemporal activity patterns of two small‐bodied benthivorous fishes, gudgeon (Gobio gobio) and stone loach (Barbatula barbatula), and of benthic invertebrates in a small temperate stream during three 24‐h field experiments. By applying a novel method of field video observation, we monitored the spatiotemporal foraging behaviour of the fish in their natural environment. A parallel analysis of invertebrate mesohabitat use by means of small area Hess sampling allowed a direct estimation of habitat overlap at a pool–riffle scale. 3. Gudgeon showed a dominant spatial activity pattern preferring pools at all times of day, whereas stone loach used both mesohabitats but with a distinct temporal (nocturnal) activity pattern. The patterns of residence were not identical with those of active foraging. Invertebrate community composition differed significantly between mesohabitats but not between times of day. More than half of the total dissimilarity between pools and riffles was accounted for by six invertebrate taxa. Five of these were subject to higher fish predation in pools than in riffles. The total prey consumption of the two fish species together in pools was about three times as high as in riffles. Trophic niche breadth of stone loach and thus its predation range was broader than that of gudgeon. 4. These results indicate that the potential predation risk for stream invertebrates depends on the combination of spatial and temporal patterns of both predator and prey. Given the distinct differences in predation risk found between pools and riffles, we conclude that spatial heterogeneity at the mesohabitat scale can influence mechanisms and consequences of selective predation. We also suggest that the analysis of spatiotemporal predator–prey relationships should not be based on the premise that the main residence habitat and active foraging habitat of a predator are identical.