Hydraulic requirements of stream communities: a case study on invertebrates

1. We relate invertebrate assemblages to direct measurements of near‐bed hydraulic conditions that integrate the complex three‐dimensional structure of flow close to the bottom. 2. We sampled invertebrate taxa from a Mediterranean River along a spatial gradient of increasing shear stress in two seasons (spring and autumn) with different hydrological conditions. We used a recently described ordination technique, Outlying Mean Index (OMI) analysis, to study the response of stream invertebrates to near‐bed hydraulic parameters. 3. The distribution of nearly 70% of the taxa collected was significantly related to the hydraulic parameters assessed. In both seasons, shear stress and Froude number were the most important hydraulic parameters whereas substratum particle size and bed roughness had less influence. Most of the 31 taxa collected in both seasons had a higher OMI (an index showing the deviation between the mean environmental conditions used by a taxon and the mean environmental conditions used by a theoretical taxon uniformly distributed across the studied gradient) in autumn (when flow was greater) and were found in samples with high shear stress and high Froude number. This suggests that benthic invertebrates changed their preferences according to flow conditions. 4. Taxon richness declined with increased shear stress during lower flow in spring. Finally, and agreeing with previous results, the proportion of filter feeders and collector‐gatherers was inversely related to shear stress. 5. Our results are a first step towards better habitat suitability models that could inform management decisions.     

When is stream invertebrate drift catastrophic? The role of hydraulics and sediment transport in initiating drift during flood events

1. The term ‘catastrophic drift’ is used to describe the large‐scale displacement of invertebrates that occurs during periods of increased river discharge. However, the physical processes that lead to animals entering the water column at such times remain poorly understood. Specifically, the hypothesis that the movement of bed sediments during floods triggers a large increase in drift has lacked a rigorous field test. 2. Using a portable flume, the hydraulic conditions and rates of bedload transport associated with small, frequent floods were created in situ within a reach of a gravel bed river. Experiments focussed on the patches of fine sediment which are the dominant source of bed material transported during small floods. The flume produced near bed velocities of up to 2 m s^?1 over the patches, increasing shear stress, initiating sediment transport and causing invertebrates to enter the drift. 3. The total number of individuals lost from the bed, as well as the taxonomic composition of the drift, were influenced strongly by shear stress and bedload. The rate of loss from the bed was low at shear stresses <9 dynes cm^?2 (0–4 individuals min^?1 from the 0.5 m² flume bed area). Once shear stress exceeded 9 dynes cm^?2, the threshold that resulted in consistent bedload transport from the patches, the rate of loss of animals increased to a maximum of 56 individuals min^?1. When bedload transport rates were at their highest, the taxonomic composition of the drift was more similar to the benthos than it was to the drift observed when bed material was stable. 4. Absolute rates of bedload transport created by the manipulations were extremely low (<7 g m^?1 s^?1) and typical of those measured during small, frequent floods. Events of this magnitude do not break up the armour layer across the reach as a whole and so exposed patches of fine sediment are the principal source of bedload material. Consequently, discharge events not considered as disturbances in geomorphic terms may initiate frequent episodes of so‐called ‘catastrophic drift’ from patches of stream bed.     

Quantifying fluid and bed dynamics for characterizing benthic physical habitat in large rivers

Sturgeon use benthic habitats in and adjacent to main channels where environmental conditions can include bedload sediment transport and high near‐bed flow velocities. Bed velocity measurements obtained with acoustic Doppler instruments provide a means to assess the concentration and velocity of sediment moving near the streambed, and are thus indicative of the bedload sediment transport rate, the near‐bed flow velocity, and the stability of the substrate. Acoustic assessments of benthic conditions in the Missouri River were conducted at scales ranging from the stream reach to individual bedforms. Reach‐scale results show that spatially‐averaged bed velocities in excess of 0.5 m s⁻¹ frequently occur in the navigation channel. At the local scale, bed velocities are highest near bedform crests, and lowest in the troughs. Low‐velocity zones can persist in areas with extremely high mean bed velocities. Use of these low‐velocity zones may allow sturgeon to make use of portions of the channel where the average conditions near the bed are severe. To obtain bed velocity measurements of the highest possible quality, it is necessary to extract bottom‐track and GPS velocity information from the raw ADCP data files on a ping‐by‐ping basis. However, bed velocity measured from a point can also be estimated using a simplified method that is more easily implemented in the context of routine monitoring. The method requires only the transect distance and direction data displayed in standard ADCP data‐logging software. Bed velocity estimates obtained using this method are usually within 5–10% of estimates obtained from ping‐by‐ping processing.     

Macroinvertebrate community structure along gradients of hydraulic and sedimentary conditions in a large gravel-bed river

1. The spatial distribution of macroinvertebrate species was examined in relation to hydraulic and sedimentary conditions in a large gravel‐bed river, the Fraser River, Canada. Mean annual discharge in the Fraser River is 2900 m3 s?1 and annual flood discharge, due to snowmelt in May and June, averages 8760 m3 s?1.
2. Invertebrates were sampled from four water depths (0.2, 0.5, 1.5, 3.0 m) at various levels of discharge that together captured the spatial and temporal variability of the physical habitat. Several hydraulic (near‐bed shear velocity, Boundary Reynolds number, turbulence intensity, depth‐averaged velocity, Froude number, Reynolds number) and substratum variables (mean grain size, Trask's sorting coefficient, Nikuradse's roughness, percentage of fine sediment, and Shields entrainment function) were measured for each sample of macroinvertebrates. Concentrations of fine and coarse particulate organic matter were also assessed.
3. The physical habitat was characterized by a major gradient of hydraulic conditions that corresponded positively with increasing water depth and accounted for 52% of the total variation in the habitat data. Substratum conditions and the concentration of organic matter explained 24% of the total variation in the habitat data.
4. The distribution of invertebrates was correlated significantly with hydraulic variables and suggests that hydraulic conditions represent a major physical gradient along which the benthic community is organized. The distribution of organic matter and substratum texture were also important for some species. The spatial distribution of most species reflected morphological and trophic suitability to particular habitat conditions.
5. Hydraulic stress associated with foraging and maintaining position, as well as organic matter retention in coarse substrata, are probable mechanisms affecting the spatial distribution of macroinvertebrates.     

The relationship between sediment mobilisation and the entry of Baetis mayflies into the water column in a laboratory flume

Mass bedload movement is thought to play a key role in initiating stream invertebrate drift during extreme flood events. However, little is known of the importance of the shear of invertebrates from stone surfaces relative to their entrainment along with bed material at different discharges. In particular, it is unclear whether so-called catastrophic drift only occurs once mass bedload movement, and hence entrainment of invertebrates, occurs. We investigated the relationship between the mobilisation and transport of bed sediments and the entry of Baetis mayflies into the water column in a laboratory flume. Experiments quantified the percentage of Baetis drifting at a range of discharges that mobilised between 0 and 95% of the flume-bed sediments. Control experiments quantified drift losses from sediment fixed to the bed of the flume, such that sediments were immobile even at the highest discharges. Drift losses increased with increasing discharge and velocity in the flume. Sediment mobility contributed significantly to drift (ANCOVA, p < 0.001), with consistently greater drift losses in mobile sediment experiments than in those with fixed sediment. The discharge which resulted in a loss of 100% of Baetis from the mobile sediment bed (discharge 30 l s^–1) resulted in a loss of approximately 50% of individuals from the fixed bed. Results indicate that once bed sediments are mobilised, entry of Baetis into the drift is greater than expected from the shear of animals from stone surfaces alone. Thus, entrainment of animals along with sediment contributes significantly to drift at high flows. This implies that differences in bed stability between sites or streams, or temporal changes in sediment characteristics within a site, could influence patterns of drift.     

Subaqueous hydrochory: open‐channel hydraulic modelling of non‐buoyant seed movement

1. Subaqueous transport may be a significant dispersal and migration mechanism of non‐buoyant seeds of aquatic and riparian plants, and also secondary transport of seeds once they have lost buoyancy, but the efficiency of this difficult to observe process is largely unexamined. This study uses hydraulic modelling to establish the discharges that move the non‐buoyant seeds of Hymenocallis coronaria as bedload or suspended load; uses stream gauge data to examine the frequency of effective discharges from late June to late September, the seed maturation and germination period; and the potential transport distance of the seeds. 2. The results show that the majority of non‐buoyant seeds of H. coronaria can be transported as bedload through entire modelled stream reaches of lengths 10.8, 18 and 14.4 km with the 0.5 year return interval flow. Bedload apparently has the ability to move seeds over great distances, and may be a substantial factor determining the genetic structure, demography and dynamics of populations and communities. However, prolonged movement of non‐buoyant seeds in suspension appears to be quite rare. 3. Although insect mediated pollination and biochory occur concurrently with bedload transport, bedload transport alone may be sufficient to account for the established gene flow rate of H. coronaria. The potential transport distance of many of the seeds exceed that between populations, and migration may occur more frequently than the species’ generation time. 4. This is the first known study to use open‐channel hydraulic modelling and sediment transport analysis to determine the effectiveness of non‐buoyant seed transport. This method of analysis shows promise for application in other contexts, and especially where flow management is a critical issue for maintenance of rare species.     

Effects of land cover on sediment regime and fish assemblage structure in four southern Appalachian streams

SUMMARY 1. We examined the relationship between catchment land cover, sediment regime and fish assemblage structure in four small streams in the upper Little Tennessee River basin of North Carolina. Study streams drained similar sized catchments (17–31 km²) with different fractions of non-forested land cover. Non-forested land cover was <3% in two 'reference' streams, whereas it was 13 and 22% in two 'disturbed' streams. Land cover data were compared with sediment transport data (suspended and bedload), benthic habitat data (embeddedness, substratum composition and coverage of fines) and fishes collected in autumn 1997.
2. Suspended sediment concentration was significantly higher in disturbed streams during both baseflow and stormflow. During baseflow disturbed streams nearly always exceeded 10 nephelometric turbidity units (NTU), whereas reference streams never exceeded this threshold. The difference in suspended sediment concentration between reference and disturbed streams was more consistent at baseflow than at stormflow. Therefore, baseflow turbidity may be a useful indicator of potential stream degradation.
3. Disturbed sites had five- to nine-fold more bedload transport than reference sites. Both embeddedness and streambed instability increased with increasing non-forested land cover.
4. Relative abundance of fishes requiring clean cobble/gravel substratum for spawning was lower in disturbed streams, whereas relative abundance of mound-building cyprinids, their nest associates and fishes that excavate nests in soft sediments (centrarchids) was higher. Relative abundance of fishes spawning in benthic crevices and gravel (BC + G) declined as the proportion of non-forested land cover increased. This study supports growing evidence that human-induced sedimentation alters stream fish assemblages.     

A novel biofilm channel for evaluating the adhesion of diatoms to non-biocidal coatings

Laboratory assessment of the adhesion of diatoms to non-toxic fouling-release coatings has tended to focus on single cells rather than the more complex state of a biofilm. A novel culture system based on open channel flow with adjustable bed shear stress values (0–2.4?Pa) has been used to produce biofilms of Navicula incerta. Biofilm development on glass and polydimethylsiloxane elastomer (PDMSe) showed a biphasic relationship with bed shear stress, which was characterised by regions of biofilm stability and instability reflecting cohesion between cells relative to the adhesion to the substratum. On glass, a critical shear stress of 1.3–1.4?Pa prevented biofilm development, whereas on PDMS, biofilms continued to grow at 2.4?Pa. Studies of diatom biofilms cultured on zwitterionic coatings using a bed shear stress of 0.54?Pa showed lower biomass production and adhesion strength on poly(sulfobetaine methacrylate) compared to poly(carboxybetaine methacrylate). The dynamic biofilm approach provides additional information to supplement short duration laboratory evaluations.     

Non-Additive Increases in Sediment Stability Are Generated by Macroinvertebrate Species Interactions in Laboratory Streams

Previous studies have shown that biological structures such as plant roots can have large impacts on landscape morphodynamics, and that physical models that do not incorporate biology can generate qualitatively incorrect predictions of sediment transport. However, work to date has focused almost entirely on the impacts of single, usually dominant, species. Here we ask whether multiple, coexisting species of hydropsychid caddisfly larvae have different impacts on sediment mobility compared to single-species systems due to competitive interactions and niche differences. We manipulated the presence of two common species of net-spinning caddisfly (Ceratopsyche oslari, Arctopsyche californica) in laboratory mesocosms and measured how their silk filtration nets influence the critical shear stress required to initiate sediment grain motion when they were in monoculture versus polyculture. We found that critical shear stress increases non-additively in polycultures where species were allowed to interact. Critical shear stress was 26% higher in multi-species assemblages compared to the average single-species monoculture, and 21% greater than levels of stability achieved by the species having the largest impact on sediment motion in monoculture. Supplementary behavioral experiments suggest the non-additive increase in critical shear stress may have occurred as competition among species led to shifts in the spatial distribution of the two populations and complementary habitat use. To explore the implications of these results for field conditions, we used results from the laboratory study to parameterize a common model of sediment transport. We then used this model to estimate potential bed movement in a natural stream for which we had measurements of channel geometry, grain size, and daily discharge. Although this extrapolation is speculative, it illustrates that multi-species impacts could be sufficiently large to reduce bedload sediment flux over annual time scales in streams where multiple species of caddisfly are present.     

Complex hydromorphology of meanders can support benthic invertebrate diversity in rivers

In freshwater environments, high biodiversity is commonly associated with habitat heterogeneity. River bends and meanders are particularly complex morphodynamic elements of watercourses. However, the specific spatio-temporal interactions between hydromorphology and the resident biota have scarcely been studied. This article reviews the relationships between hydraulic processes, and morphological units that are typical for meanders, and analyzes the concomitant spatial and temporal dynamics of habitats suitable for aquatic invertebrates. Flow in river bends is characterized by significant cross-stream velocities, which modify primary flow patterns, and create helical flow trajectories. Consequently, boundary shear stresses at the river-bed are altered, so that complex erosion, transport, and accumulation processes characteristically shape bed and bank morphology. The diversity of substrate types and complex bathymetry in meanders provide a large variety of habitat conditions for benthic invertebrates within a relatively small spatial domain, which are connected via hydraulic pathways. Periodic reversal of hydro-morphological processes between low and high flow, and seasonal growth of aquatic macrophytes creates spatio-temporal dynamics at the meso- and microhabitat scales. Such habitat dynamics increases benthic invertebrate diversity to the extent it is consistent with spatio-temporal scales of invertebrate mobility and life cycle. Furthermore, the presence of flow refugia, and hydraulic dead zones in meanders is essential to sustain species richness. This study concludes that meanders are highly complex morphodynamic elements that exhibit several self-regulating principles supporting invertebrate diversity and resilience in fluvial ecosystems.     

LINKING BENTHIC ALGAL BIOMASS TO STREAM SUBSTRATUM TOPOGRAPHY1

The physical properties of substrata significantly influence benthic algal development. We explored the relationships among substratum surface texture and orientation with epilithic microphytobenthic biomass accumulation at the whole‐substratum and micrometer scales. Unglazed clay tiles set at three orientations (horizontal, vertical, and 45°), and six substrata of varying surface roughness were deployed in a prairie stream for 3 weeks. Substrata were analyzed for loosely attached, adnate, and total benthic algal biomass as chl a, and confocal laser scanning microscopy was used to measure substrata microtopography (i.e., roughness, microscale slope angles, and three‐dimensional surface area). At the whole‐substratum level, vertical substrata collected significantly (P < 0.05) less algal biomass, averaging 34% and 36% less than horizontal and 45° substrata, respectively. Benthic algal biomass was also significantly less on smoother surfaces; glass averaged 29% less biomass than stream rocks. At the microscale level, benthic algal biomass was the greatest at intermediate values, peaking at a mean roughness of approximately 17 μm, a mean microscale slope of 50°, and a projected/areal surface area ratio of 2:1. The proportion of adnate algae increased with surface roughness (26% and 67% for glass and brick, respectively), suggesting that substratum type changes the efficiency of algal removal by brushing. Individual substrata and microsubstrata characteristics can have a strong effect on benthic algae development and potentially affect reach scale algal variability as mediated by geomorphology.     

Stream geomorphology regulates the effects on periphyton of ecosystem engineering and nutrient enrichment by Pacific salmon

1. Pacific salmon (Oncorhynchus spp.) returning to streams deliver substantial quantities of nutrients (nitrogen and phosphorus) that may stimulate primary production. Salmon can also affect the phytobenthos negatively via physical disturbance during nest excavation, a process that may counteract the positive effects of salmon‐derived nutrients on benthic algae. The ability of salmon to disturb benthic habitats may be a function of substratum particle size, and therefore, the geomorphology of streams could determine the net effect of salmon on benthic communities. 2. Based on surveys of 17 streams in southwest Alaska before the salmon run and during peak salmon density, we identified size thresholds for the disturbance of substratum particles by salmon and classified particles as vulnerable (<60 mm B‐axis), invulnerable (>110 mm) or transitional (61–110 mm). At the scale of individual rocks, algal biomass on vulnerable substrata decreased at peak spawning (relative to values before the run) as a power function of salmon density; transitional and invulnerable substrata showed no quantifiable pattern. However, invulnerable substrata in streams with more than 0.11 salmon m^?2 showed net algal accrual, or relatively smaller declines in algal biomass, than vulnerable substrata, indicating that large rocks provide refuge for benthic algae from salmon disturbance. 3. We expected that streams with proportionally larger rocks would respond positively to salmon at the whole‐stream scale, after accounting for the relative abundance of rocks of different sizes within streams. Invulnerable rocks made up only 0–12% of the total substratum particle size distribution in salmon‐bearing streams, however, and algal accrual on invulnerable substrata did not outweigh the strong disturbance effects on the more spatially extensive vulnerable substrata. The change in whole‐stream benthic algal biomass among streams was negatively related to salmon density. 4. Stable isotopes of nitrogen (δ¹⁵N) were used to track nutrients from salmon into benthic biota. Periphyton δ¹⁵N on rocks of all size classes was higher at peak salmon spawning than before the salmon run, indicating the uptake of salmon‐derived nitrogen. Peak δ¹⁵N values were positively related to salmon abundance and followed a two‐isotope mixing relationship. The per cent of N from salmon in periphyton was also related to salmon density and was best explained by a saturating relationship. Spring δ¹⁵N was unrelated to salmon returns in the previous year, suggesting little annual carryover of salmon nutrients.     

Influences of Woody Debris on Flow Patterns and Channel Morphology in a Low Energy,Sand‐Bed Stream Reach

In lowland areas, such as the glacial landscapes of eastern Germany, sand‐bed streams are the most common stream type. They have low gradients and their hydrological regime is often subdued due to the frequent interruption by lakes. Very few is known about the influence of woody debris in these streams, since nearly all previous studies are from high‐gradient conditions, where streams have coarse bed sediments and harsh hydrological regimes. The research objectives of this study were first to assess the quasi‐natural quantity and quality of wood in a lowland sand‐bed stream and second to understand the influence of wood on the channel morphology and the flow patterns at base‐flow. The three‐dimensional stream bed relief was surveyed by electronic distance measurement. The position and the size of large woody debris was assessed by close‐up photography. An acoustic Doppler velocimeter was used to record the patterns of flow velocity and turbulence. Overlay and analysis of the spatial data was done using a Geographic Information System. The standing stock of wood was 1.9 m³ and 39 woody elements per 100 m² of stream bed. The flow pattern was clearly controlled by the wood. Woody elements elevated above the stream bed deflected flow and locally caused strong secondary current, high turbulence, and scour of the stream bed at baseflow. Wood resting directly on the stream bed, which contributed the majority of the wood inside the bank‐full channel, determined the roughness of the stream bed. Near‐bed flow patterns observed were isolated roughness flow and wake interference flow, which was registered inside the accumulations of wood. 68% of the stream bed had shear stress above critical. Hence, the secondary morphological structures of the sand‐bed were controlled at base‐flow by the flow which was determined by the woody debris distribution.     

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

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

Microdistribution of benthic invertebrates in a rocky mountain (U.S.A.) stream

A study of the benthic invertebrate community inhabiting a small, foothill trout stream in the Rocky Mountains of Idaho was conducted over a two-year period. Monthly Hess samples and short-term experiments using substratum-filled trays were used to describe the spatial dispersion of the benthos and to examine the response of invertebrate populations to substratum and current. A method was devised for measuring available surface area which involved coating individual stones with latex and measuring the area of the print resulting from inking the impression left on the latex mold.The dispersion of all populations was clumped throughout the year. Alteration of the cross-sectional pattern of current velocity and stream bed composition changed the pattern of distribution but not the extent of clumping. Collections made in areas of depositing and eroding substrata revealed a more diverse fauna in the latter. Most groups of organisms found in the riffle were scarcer in the pools or absent from them. The pool fauna contained no important additions over those found in the riffles.After a year's study of invertebrate populations in an otherwise undisturbed riffle, the substratum was altered and the flow made more uniform; an increase in the abundance of most of the benthic invertebrates followed. No single factor was responsible for the increase, but the change in substratum size and degree of compaction accounted for most of the change. Interpretation of the results was aided by findings from experiments using substratum-filled trays.Two series of stream experiments using the trays were conducted: one to test the relative importance of current and substratum and the other to test the effect of particle size on the distribution of the benthic fauna. In the first series, placement of trays of stones in a pool resulted in an increase in numbers of some but not all of the invertebrates over numbers usually occurring in the pool. Trays filled with stones and placed in a riffle supported fewer animals than found on the adjacent stream bed but more than in the pool. Variations are attributed to differences in current velocity and amounts of imported organic and inorganic debris. Three different relationships of population numbers to current velocity were found for different members of the community (direct, indirect, and parabolic) over the range of 10 to 60 cm/sec. The second series of experiments consisted of two sets of trays filled with stones of medium or large pebbles, respectively. Nine taxa, as well as all of the combined taxa, showed a preference for trays of small stones over the natural stream bed. A few taxa were noticeably more abundant on the small substratum than on the large but most of the fauna showed only slight increases in numbers or remained constant on the two substrata. Only three taxa showed a direct relation of numbers to total surface area presented by the stones.Number and kinds of organisms found in trays filled with a uniform size of substratum did not correspond to those taken in Hess samples from the natural stream bed. This has important implications in terms of currently recommended pollution monitoring techniques. However, it is suggested that if the substratum composition of the trays more nearly matched that of the stream, the correspondence would be much better. The results of the present study also throw considerable doubt on the adequacy of generalizations derived from earlier studies of responses to substratum size and suggest several reasons for reevaluating current ideas regarding the influence of substratum on invertebrate distribution.     

Habitat attributes associated with short‐term settlement of Ozark hellbender (Cryptobranchus alleganiensis bishopi) salamanders following translocation to the wild

1. Organisms associated with lotic systems rank among the most threatened because of global change. Although translocation is being increasingly applied as a conservation strategy, most studies have focused on survival and recruitment of individuals, and few have attempted to identify how habitat attributes influence short‐term settlement of animals during the critical post‐release period. 2. We demonstrate the application of resource selection modelling in an information theoretic framework to identify release‐site characteristics that will increase the likelihood of settlement for a fully aquatic benthic stream salamander, the Ozark hellbender (Cryptobranchus alleganiensis bishopi). We fit discrete choice models using data from 29 radio‐tagged hellbenders that were translocated to two sites in the North Fork of the White River (NFWR), Missouri (U.S.A.). We defined resource availability at two spatial scales (stream reach and home range) and quantified abiotic habitat attributes at 3181 salamander locations and 6329 random available locations collected between May 2008 and August 2009. 3. At both sites and spatial scales, a single model received substantially greater support (0.96–1.00 of total model weight) than all other models, and top‐ranked models were similar in form and predictive ability. At both spatial scales, selection was positively influenced by the presence of cobble‐boulder substratum relative to bedrock and finer substrata. We also noted a negative interactive effect between distance to the nearest substratum particle large enough to provide cover (i.e. at least one axis ≥15 cm in length) and an increase in either a direct or relative (i.e. pool, run, and riffle) measure of water velocity. 4. Collectively, salamanders released in our study selected resources indicative of long‐term benthic microhabitat stability. However, despite strong selection of cobble‐boulder substratum, 8% (282 of 3181) of captive‐reared hellbender locations occurred in bank crevices and root masses. Although several studies have reported the importance of near bed hydraulics in determining occurrence of stream macroinvertebrates, our findings are the first to indicate that spacing among cobble‐boulder substrata may be important for hellbenders. 5. To increase the likelihood of short‐term settlement of captive‐reared hellbenders in the wild, we recommend prioritising release sites where the average distance between cobble‐boulder particles within habitat patches is minimised. In general, average spacing among cobble and boulder substrata should be <1 m in habitat patches where mean benthic water velocity exceeds 0.1 m s^?1, and <0.5 m where water velocity approaches 0.30 m s^?1. Based on home range sizes of captive‐reared Ozark hellbenders, the collective extent of suitable cobble‐boulder habitat patches within release sites should approximate at least 10 m² per salamander released.     

Flow- and substratum-mediated movement by a stream insect

1. In streams subject to frequent hydrologic disturbance, the ability of benthic invertebrates to disperse within the channel is key to understanding the mechanisms of flow refugium use and population persistence. This study focuses on crawling invertebrates, the effects on movement of abiotic factors (specifically, flow near the stream bed and bed micro‐topography) and the consequences for dispersal. 2. In a large flume, we observed individual cased caddisflies, Potamophylax latipennis, moving in fully turbulent flows over a precise replica of a water‐worked surface. From maps of movement paths, we quantified crawling behaviour and entrainment, and the influence of bed micro‐topography. We manipulated discharge and tested its effect on movement, linear displacement and areal dispersal. The highest discharge treatment was a disturbance to the caddis; the lowest discharge was not. Crawling behaviours were used to parameterise random walk models and estimate population dispersal, and to test the effects of abiotic factors on movement. 3. Bed micro‐topography influenced crawling in several ways. Caddis spent most of their time at the junctions between proud particles and the adjacent plane bed. The frequency distribution of turn angles was bimodal, with modal values approximating the angle required to travel around median‐sized particles. Larvae generally crawled downstream, but crawling direction relative to the flow was skewed by bed micro‐topography and was not directly downstream, unlike drift. 4. Caddis crawled for most of the time and discharge affected almost every aspect of their movement. As discharge increased, caddis crawled less often, more slowly and over shorter distances; they also became entrained more frequently and over greater distances. With increased discharge, caddis spent proportionately less time at the junctions between proud particles and the adjacent plane bed, and more time on the tops and sides of proud clasts. This is curious as most entrainment occurred from the tops and sides of clasts and entrainment is generally considered to be disadvantageous during disturbances. 5. Linear displacement (drift and entrainment combined) was downstream, but the relation between total displacement and discharge was complex. Total displacement decreased at intermediate discharge as crawling decreased, but increased at high discharge as entrainment and drift played a greater role in movement. 6. Within‐stream dispersal via crawling contained elements of both a correlated random walk (we observed directional persistence in turn angles) and a biased random walk (we observed downstream bias in move direction angles) and was best described as a biased correlated random walk. Dispersal was inversely related to discharge, suggesting that the ability of P. latipennis to crawl into flow refugia on the streambed is reduced at high flow.     

Physicochemical stream bed characteristics and recruitment of the freshwater pearl mussel (Margaritifera margaritifera)

1. The freshwater pearl mussel (Margaritifera margaritifera) is endangered and of conservation importance. We used its survival/mortality during the critical post‐parasitic phase as a biological indicator for the habitat quality of the stream substratum. 2. We established and tested biological, physical and chemical methods of assessing the stream bed in 26 streams from seven European countries. We analysed penetration resistance, texture, the concentrations and ratios of C, N, S, P, Fe, Mn in fine material <100 μm, and redox, pH and electric conductivity at the surface and at 5 and 10 cm into the substratum. 3. Sites with high stream bed quality (promoting pearl mussel populations with good juvenile recruitment) had coarser and better sorted substrata with significantly lower quantities of fines, and a higher Mn concentration in the fines, than poor quality sites. Redox potential (Eh) at sites without recruitment differed markedly between the free‐flowing water at the surface and at 5 and 10 cm in the bed, whereas no differences were detectable at good quality sites. This was also true of electric conductivity and, to a lesser extent, pH. The stream bed at sites lacking pearl mussel recruitment had a more variable and higher penetration resistance, indicating clogging of the interstitial macropore system by the deposition of mud and compaction of the stream bed. 4. Our results show that habitat quality for pearl mussels depends strongly on the exchange between the surface and the interstices, which is governed by physicochemical characteristics of the stream substratum. Combined measurements of penetration resistance, depth gradients of Eh and texture were most suitable for assessing stream bed quality, while water chemistry was insufficient because of the decoupling of interstitial and free‐flowing water at poor quality sites.     

Substratum stability associated with the riverine macrophyte Justicia americana

1. Patches of stable substratum in streams may be important refugia for benthic organisms during scouring floods. Streambed stone stability, packing and embeddedness were assessed within and adjacent to beds of the macrophyte Justicia americana in five Alabama streams. 2. The force needed to dislodge stones and embeddedness was about two times lower outside Justicia beds than within them. Significant positive correlations between stone stability and (i) degree of embeddedness, and (ii) the abundance of binding rhizomes and the presence of attached roots indicate that Justicia may physically modify the local streambed, indirectly enhancing substratum stability and reducing flow, thereby increasing sand deposition. 3. Despite higher stability (i.e. physical refugia during bed‐moving spates) within Justicia beds, the abundance of epilithic plants (moss and Podostemum ceratophyllum) and pleurocerid snails (Elimia spp.) was similar both inside and outside the macrophyte beds. Several physical characteristics within macrophyte beds, such as low light, reduced current and increased sand intrusion, may create suboptimal conditions for benthic organisms in these habitats. 4. Additional work is needed to determine if Justicia biogenically enhances substratum stability or if its presence merely reflects patches of stable substratum within the streambed. Regardless of the mechanism, there is an association between Justicia beds and streambed characteristics.     

Stream channel geomorphology influences mussel abundance in southern Appalachian streams, U.S.A.

1. Studies of North American streams have shown that hydraulic parameters and stream geomorphology can explain unionid mussel abundance at both the reach and catchment scale. However, few studies have examined applicability of hydrogeomorphic variables across broader spatial scales, such as across whole catchments, or have elucidated conditions under which spates can affect mussel populations in streams. 2. We quantified freshwater mussel abundance and species richness and their physical habitat at 24 sites in eight streams in southern Appalachian catchments in 2000 and 2001. In addition, we modelled site‐specific hydraulic parameters during summer baseflow and bankfull stages to estimate high‐ and low‐discharge conditions, respectively. 3. Mussel abundance was related to stream geomorphology, whereas richness was related to stream size. Baseflow habitat parameters explained only minor variation in abundance or richness, and both measures were highly correlated with mean current velocity or stream size. Bankfull shear stress composed a relatively low proportion of overall mussel habitat variability, but it accounted for significant variation in abundance and richness. 4. Mussel abundance was highly variable at sites subject to low‐shear stress during spates, whereas abundance always was low at sites subject to high‐shear stress. These data suggest that habitat conditions during floods, rather than those at summer baseflow, limit the abundance of mussels in Appalachian streams. These data also suggest that mussel abundance and assemblage structure may be sensitive to any changes in channel geomorphology and hydraulic conditions that might result from land use in the catchment.