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     

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.     

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.     

Spatial and Temporal Variation of Macroinvertebrate Drift in Two Neotropical Streams1

The detection of spatial variation in macroinvertebrate drift depends on the spatial scale of investigation in streams of the La Selva Biological Station, Costa Rica. Drift samples were taken in a spatially nested design, with two streams, two reaches per stream, two riffles per reach, and four replicate samples per riffle. Drift showed little variation among streams, but varied significantly at the scales of reach and riffle, with variation among samples also high. In addition, sampling took place at two temporal scales: diel and at two different periods that differed in rainfall conditions. Drift diel periodicity was a clear pattern, while only density of individuals varied among sampling periods. This is the first study of macroinvertebrate drift at multiple spatial scales, despite the recognition that multi‐scale studies are essential for a more complete understanding of community patterns and processes.     

The spatial structure of variation in salamander survival,body condition and morphology in a headwater stream network

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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.     

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.     

Covariation of stream community structure and biomass of algae, invertebrates and fish with forest cover at multiple spatial scales

1. To evaluate the spatial extent of the effects of forest cover on stream ecosystems, we measured algae, invertebrate, and fish biomass and invertebrate and fish community structure in 38 small first- to third-order streams in the National Capital Region of Canada along with forest cover at different spatial scales.
2. We considered 55 spatial scales of forest cover including several buffer widths (doubling 10–320 m) and lengths (doubling 10–1280 m, entire riparian distance upstream from sampling area) and entire catchments to determine which spatial scale maximized the correlation with biomass and metrics of community structure.
3. The proportion of variability in biomass and structural metrics explained by forest cover generally increased with increasing scale, suggesting that catchment-wide disturbances are the most influential determinants of benthic and fish communities.
4. Catchment forest cover explained more variation in algal (adjusted r ²   =   0.54), invertebrate (adjusted r ²   =   0.51) and fish (adjusted r ²   =   0.33) biomass than structural metrics of invertebrates and fish (adjusted r ²   =   0.08–0.27).
5. Analyses of the partial effects of forest cover at three scales (reach, riparian and the entire catchment) on biomass and community structure metrics identified catchment and reach scales as being most influential and never detected a significant partial effect of forest cover at the riparian scale.
6. These results suggest that maintenance or protection of reach and riparian buffers alone will not sufficiently protect stream function and structure from catchment-wide impacts.     

Identifying the scales of variability in stream macroinvertebrate abundance, functional composition and assemblage structure

1. Many natural ecosystems are heterogeneous at scales ranging from microhabitats to landscapes. Running waters are no exception in this regard, and their environmental heterogeneity is reflected in the distribution and abundance of stream organisms across multiple spatial scales. 2. We studied patchiness in benthic macroinvertebrate abundance and functional feeding group (FFG) composition at three spatial scales in a boreal river system. Our sampling design incorporated a set of fully nested scales, with three tributaries, two stream sections (orders) within each tributary, three riffles within each section and ten benthic samples in each riffle. 3. According to nested anova s, most of the variation in total macroinvertebrate abundance, abundances of FFGs, and number of taxa was accounted for by the among‐riffle and among‐sample scales. Such small‐scale variability reflected similar patterns of variation in in‐stream variables (moss cover, particle size, current velocity and depth). Scraper abundance, however, varied most at the scale of stream sections, probably mirroring variation in canopy cover. 4. Tributaries and stream sections within tributaries differed significantly in the structure and FFG composition of the macroinvertebrate assemblages. Furthermore, riffles in headwater (second order) sections were more variable than those in higher order (third order) sections. 5. Stream biomonitoring programs should consider this kind of scale‐dependent variability in assemblage characteristics because: (i) small‐scale variability in abundance suggests that a few replicate samples are not enough to capture macroinvertebrate assemblage variability present at a site, and (ii) riffles from the same stream may support widely differing benthic assemblages.     

Taxonomic and functional homogenisation of macroinvertebrate communities in recently intermittent Alpine watercourses

1. Mountain streams in southwestern European Alps are currently shifting from perennial to intermittent flow due to the combined effects of climate change and local anthropogenic pressures. Given that flow intermittency is a recently documented phenomenon in the Alps, only scattered studies have investigated functional and taxonomical diversity of benthic invertebrate communities in recently intermittent Alpine streams.
2. We used a hierarchical sampling design to investigate patterns in taxonomic and functional diversity of benthic invertebrate communities in 13 recently intermittent Alpine streams in north-west Italy. in April 2017, we sampled benthic communities in two reaches of each stream with different hydrological conditions: a control reach, with permanent flow; and an intermittent reach, which recently experienced non-flow periods in summer.
3. We tested for the response of taxonomic richness at multiple spatial scales by partitioning total diversity into the average richness of local communities and the richness due to variation among local communities both within and among reaches. By partitioning total diversity (γ) into its local (α) and turnover (β) components we showed a decrease in local and regional species richness both within and among reaches, whereas variation among communities was significantly lower in intermittent reaches at the reach scale only.
4. The analysis of multidimensional trait space of macroinvertebrate communities in reaches with different hydrological conditions revealed a significant reduction of functional diversity, dispersion, and evenness in intermittent reaches. There was trait overdispersion in intermittent reaches, as these hosted both typical Alpine taxa and organisms adapted to flow intermittency. In particular, we observed the replacement of taxa with aquatic respiration and those preferring medium- to fast-flowing oligotrophic waters by taxa adapted to lentic habitats, air breathing and with larval dormancy phases.
5. These results indicate that recent flow intermittency has caused drastic changes in benthic invertebrate communities in Alpine streams. Our work highlights the importance of integrating taxonomic and functional diversity to thoroughly assess the impacts of flow intermittency.

     

Stream communities across a rural–urban landscape gradient

Rapid urbanization throughout the world is expected to cause extensive loss of biodiversity in the upcoming decades. Disturbances associated with urbanization frequently operate over multiple spatial scales such that local species extirpations have been attributed both to localized habitat degradation and to regional changes in land use. Urbanization also may shape stream communities by restricting species dispersal within and among stream reaches. In this patch-dynamics view, anthropogenic disturbances and isolation jointly reduce stream biodiversity in urbanizing landscapes. We evaluated predictions of stream invertebrate community composition and abundance based on variation in environmental conditions at five distinct spatial scales: stream habitats, reaches, riparian corridors and watersheds and their spatial location within the larger three-river basin. Despite strong associations between biodiversity loss and human density in this study, local stream habitat and stream reach conditions were poor predictors of community patterns. Instead, local community diversity and abundance were more accurately predicted by riparian vegetation and watershed landscape structure. Spatial coordinates associated with instream distances provided better predictions of stream communities than any of the environmental data sets. Together, results suggest that urbanization in the study region was associated with reduced stream invertebrate diversity through the alteration of landscape vegetation structure and patch connectivity. These findings suggest that maintaining and restoring watershed vegetation corridors in urban landscapes will aid efforts to conserve freshwater biodiversity.     

Beta diversity of stream diatoms at two hierarchical spatial scales: implications for biomonitoring

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Testing for temporal coherence across spatial extents: the roles of climate and local factors in regulating stream macroinvertebrate community dynamics

Temporal coherence or spatial synchrony refers to the tendency of population, community or ecosystem dynamics to behave similarly among locations through time as a result of spatially‐correlated environmental stochasticity (Moran effect), dispersal or trophic interactions. While terrestrial studies have treated synchrony mainly as a population‐level concept, the majority of freshwater studies have focused on community‐level patterns, particularly in lake planktonic communities. We used spatially and temporally hierarchical data on benthic stream invertebrates across six years, with three seasonal samples a year, in 11 boreal streams to assess temporal coherence at three spatial extents: 1) among regions (watersheds), 2) among streams within a region, and 3) among riffles within a stream, using the average of correlation coefficients for stream/riffle pairs across years. Our results revealed the primacy of strongly synchronized climatic factors (precipitation, air temperature) in inducing temporal coherence of macroinvertebrate assemblages across geographically distinct sites (i.e. Moran effect). Coherence tended to decrease with increasing spatial extent, but positive coherence was detected for most biological variables even at the largest extent (about 350 km). The generally high level of coherence reflected the strong seasonality of boreal freshwater communities. A hydrologically exceptional year enhanced the synchrony of biological variables, particularly total macroinvertebrate abundance. Regionally low precipitation in that year led to a substantial decrease in benthic densities across a broad spatial extent, followed by a rapid post‐drought recovery. Coherence at the among‐riffle (within‐stream) extent was lower than expected, implying that local‐scale habitat filters determine community dynamics at smaller spatial extents. Thus, temporal coherence of stream benthic communities appears to be controlled by partly different processes at different spatial scales.     

Habitat characteristics and trophic structure of benthic macroinvertebrates in a forested headwater stream

Three channel reaches with different habitat characteristics were selected to test the variability in community structure of benthic macroinvertebrates by comparing the relative abundance of functional feeding groups among the reaches. The important factors influencing the spatial and temporal organization of community structure were explored using nonmetric multidimensional scaling (NMS). The habitat characteristics in the reaches were different in terms of habitat type, hydrological factors, and substrate composition. The first headwater reach was classified as a step-pool reach with similar relative areas of riffle and pool habitats. The second mid-reach and the third down reach had greater areas of pool habitat followed by runs and riffles whose proportions were similar between the latter two reaches. The relative abundances of functional feeding groups were different among the surveyed reaches. Gammarid shredders predominated in the upper reach, and chironomid collector–gatherers and collector–filterers were in greater abundance in the two lower reaches. The proportions of gammarids were minor in the mid and downstream reaches. NMS ordination indicated that the proportion of substrates < 8 mm, discharge, and water depth mainly determined the spatial and temporal distribution of samples based on the macroinvertebrate community in the study reaches. These results suggest that different habitat characteristics result in a distinct community structure in each reach.     

Relationship between Riparian Vegetation and Stream Benthic Communities at Three Spatial Scales

We examined the influence of riparian vegetation on macroinvertebrate community structure in streams of the Upper Thames River watershed in southwestern Ontario. Thirty-three μ-basins (129–1458 ha) were used to identify land cover variables that influenced stream macroinvertebrates. Micro-basins represented the entire drainage area of study streams and were similar in stream order (first, second) and land cover (agricultural or forest; no urban). We described the structure and composition of riparian vegetation and benthic macroinvertebrate communities at the outflow reach. The nature of the land cover was quantified for the stream network buffer (30 m) and the whole μ-basin. The objective of this study was to measure the magnitude and nature of the relationship between the riparian vegetation and benthic macroinvertebrate community at the outflow reach, stream network buffer, and whole μ-basin scales. Taxon richness (including total number of Ephemeroptera, Plecoptera, and Trichoptera taxa) and Simpson’s diversity of the macroinvertebrate community all increased with increased tree cover in the riparian zone at the outflow reach scale. Simpson’s equitability was lower with greater agricultural land cover in the stream network buffer. No relationship between the macroinvertebrate community and land cover was found at the whole μ-basin scale. Analysis of the influence of land cover on stream communities within a spatial hierarchy is important for understanding the interactions of stream ecosystems with their adjacent landscapes.     

Effects of fine sediment addition and removal on stream invertebrates and fish: a reach‐scale experiment

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Do isolation and local habitat jointly limit the structure of stream invertebrate assemblages?

1. Both local and regional processes simultaneously control species assemblages depending on spatial habitat configuration. In dendritic networks like streams, the unique spatial arrangement of habitats produces various combinations of local habitat size and isolation. Stream invertebrate assemblages could therefore be controlled by different combinations of local and regional processes, depending on their location in the network. 2. Using quantile regression, we investigated how local habitat size, local environmental conditions and spatial isolation influenced variation in assemblage composition. Adult Trichoptera and benthic macroinvertebrate assemblages were represented by non‐metric multidimensional scaling (NMDS) ordination scores, as were local environmental conditions, in four headwater stream networks in New Zealand. 3. With increasing local habitat size, there was a decrease in variation in assemblage composition (NMDS scores) of both adult Trichoptera and benthic macroinvertebrates. This relationship between habitat size and assemblage variation was related to local habitat conditions at the upper limit of assemblage variability and spatial isolation at the lower limit of assemblage variability, for both adult Trichoptera and benthic assemblages, indicating joint local and regional controls on stream invertebrate assemblages. 4. The relationships between local assemblages and their neighbours, based on community similarity scores, differed between benthic macroinvertebrates and adult Trichoptera. For benthic assemblages, the larger the stream, the more similar assemblages were to neighbouring assemblages, whereas there was no consistent relationship between assemblage similarity and stream size for adult Trichoptera. This difference in structuring could be attributed to contrasting spatial influences linked to the different dispersal modes of adults and larvae. However, because adult and benthic assemblages are not independent, the influence of life stage on spatial distribution is difficult to determine (i.e. it is essentially a ‘chicken and egg’ argument). 5. Overall, our approach using quantile regression to evaluate limit responses, rather than regressions on means, has highlighted the joint importance of local habitat and spatial processes in structuring stream invertebrate assemblages. Furthermore, we have provided evidence for the importance of the spatial network arrangement and interactions between life stages and dispersal processes, in structuring stream assemblages.     

Natural‐Channel‐Design Restorations That Changed Geomorphology Have Little Effect on Macroinvertebrate Communities in Headwater Streams

Stream restorations that increase geomorphic stability can improve habitat quality, which should benefit selected species and local aquatic ecosystems. This assumption is often used to define primary restoration goals; yet, biological responses to restoration are rarely monitored or evaluated methodically. Macroinvertebrate communities were inventoried at 6 study reaches within 5 Catskill Mountain streams between 2002 and 2006 to characterize their responses to natural‐channel‐design (NCD) restoration. Although bank stability increased significantly at most restored reaches, analyses of variation showed that NCD restorations had no significant effect on 15 of 16 macroinvertebrate community metrics. Multidimensional scaling ordination indicated that communities from all reach types within a stream were much more similar to each other within any given year than they were in the same reaches across years or within any type of reach across streams. These findings indicate that source populations and watershed‐scale factors were more important to macroinvertebrate community characteristics than were changes in channel geomorphology associated with NCD restoration. Furthermore, the response of macroinvertebrates to restoration cannot always be used to infer the response of other stream biota to restoration. Thus, a broad perspective is needed to characterize and evaluate the full range of effects that restoration can have on stream ecosystems.