Reach- and catchment-scale determinants of the distribution of freshwater mussels (Bivalvia: Unionidae) in south-eastern Michigan, U.S.A.

1. We investigated the diversity and distribution of freshwater mussels at 40 sites in an agricultural catchment, the River Raisin in south‐eastern Michigan, to relate mussel assemblages and individual taxa to reach and catchment‐scale variables. Unionids were surveyed by timed searches in 100‐m reaches, and in‐stream and riparian habitat were quantified as well as flow, water chemistry and channel morphology. Land use/cover and surficial geology were determined for site subcatchments and riparian buffers. 2. Some 21 mussel species were found overall; richness ranged from 0 to 12 living species per site. From the upper to middle to lower catchment, the number of individuals, number of species, Shannon–Weaver diversity and relative abundance of intolerant unionids all declined significantly. 3. Four groupings based on overall mussel diversity and abundance were significantly related to reach‐scale habitat variables. The richest mussel assemblages were associated with sites with higher overall habitat quality, greater flow stability, less fine substratum, and lower specific conductance. 4. Stepwise multiple regressions revealed that the distribution and abundance of the total mussel assemblage, as well as the most common species, could be predicted from a combination of reach‐ and catchment‐scale variables (R² = 0.63 for total mussels, R² = 0.51–0.86 for individual species). 5. Flow stability, substratum composition and overall reach habitat quality were the most commonly identified reach‐scale variables, and measures of surficial geology were the most effective catchment‐scale variables. The spatial pattern of geology is likely to be responsible for the diversity gradient from the upper to the lower catchment. 6. Prior studies, attempting to explain mussel distributions from local habitat features alone, have found relatively weak relationships. By employing a combination of reach‐ and catchment‐scale habitat variables, this study was able to account for a substantial amount of the spatial variability in mussel distributions.     

Geomorphology and fish assemblages in a Piedmont river basin, U.S.A.

1. We investigated linkages between fishes and fluvial geomorphology in 31 wadeable streams in the Etowah River basin in northern Georgia, U.S.A. Streams were stratified into three catchment sizes of approximately 15, 50 and 100 km², and fishes and geomorphology were sampled at the reach scale (i.e. 20–40 times stream width). 2. Non‐metric multidimensional scaling (NMDS) identified 85% of the among‐site variation in fish assemblage structure and identified strong patterns in species composition across sites. Assemblages shifted from domination by centrarchids, and other pool species that spawn in fine sediments and have generalised food preferences, to darter‐cyprinid‐redhorse sucker complexes that inhabit riffles and runs, feed primarily on invertebrates, and spawn on coarser stream beds. 3. Richness and density were correlated with basin area, a measure of stream size, but species composition was best predicted (i.e. |r| between 0.60–0.82) by reach‐level geomorphic variables (stream slope, bed texture, bed mobility and tractive force) that were unrelated to stream size. Stream slope was the dominant factor controlling stream habitat. Low slope streams had smaller bed particles, more fines in riffles, lower tractive force and greater bed mobility compared with high slope streams. 4. Our results contrast with the ‘River Continuum Concept’ which argues that stream assemblages vary predictably along stream size gradients. Our findings support the ‘Process Domains Concept’, which argues that local‐scale geomorphic processes determine the stream habitat and disturbance regimes that influence stream communities.     

Beaver dams maintain fish biodiversity by increasing habitat heterogeneity throughout a low‐gradient stream network

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Hard rock versus soft bottom: the fauna associated with intertidal mussel beds on hard bottoms along the coast of Chile,and considerations on the functional role of mussel beds

The fauna associated with hard bottom mussel beds along the exposed Pacific coast of Chile was examined. The abundance of adult (>10 mm body length) purple mussels Perumytilus purpuratus varied between 32 and 75 individuals per 50 cm², and their biomass between 4.8 and 8.6 g AFDW per 50 cm² at eight sampling sites between Arica (18°S) and Chiloé (42°S). At all sampling sites, the associated fauna was dominated by suspension-feeding organisms (cirripeds, spionid and sabellid polychaetes, a small bivalve) followed by grazing peracarids and gastropods. Predators and scavengers also reached high abundances while deposit- and detritus-feeding organisms were of minor importance. The majority of organisms associated with these hard bottom mussel beds feed on resources obtained from the water column or growing on the mussels rather than on materials deposited by the mussels. This is in contrast to the fauna associated with mussel beds on soft bottoms, which comprises many species feeding on material accumulated by mussels (faeces and pseudofaeces) and deposited within the mussel bed. Many of the organisms dwelling between mussels both on hard bottoms and on soft bottoms have direct development, but organisms with pelagic development also occur abundantly within mussel beds. We propose that species with direct development are disproportionately favoured by the structurally complex habitat with diverse interstitial spaces between the mussels, which provides ample shelter for small organisms. We conclude that mussels on hard-bottoms primarily provide substratum for associated fauna while mussels on soft bottoms provide both substratum and food resources. Electronic Publication     

Effects of substratum restoration on salmonid habitat quality in a subalpine stream

Stream substratum restoration is a widely applied tool to improve spawning habitat quality for salmonid fishes. However, there is a lack of studies which comprehensively assess effects of the restoration on site, as well as on downstream habitats. Our study addressed effects at both locations and compared abiotic (analyses of texture, penetration resistance, oxygen concentration, redox, nitrite, nitrate, ammonium, pH, electric conductivity, temperature) with biotic (depth-specific macroinvertrebrate abundance and diversity, brown trout hatching success) indicators before and after excavation of the substratum in a highly colmated brown trout spawning site. Strong improvements of hyporheic water conditions (increased oxygen supply and redox potential, reduced concentrations of nitrite and ammonium) as well as ~50 % reductions of substratum compaction and fine sediment content were observed 1 day after the restoration measure. Improvements of habitat quality were still detectable 3 months after treatment. Consequently, the hatching success of Salmo trutta eggs increased from 0 % to 77 % after the restoration. Short-term decrease of macroinvertebrate abundance (from 13.1 to 3.9 macroinvertebrates/kg substratum) was observed within the hyporheic zone of the restoration site, but after 3 months, the number of taxa increased from 13 to 22 taxa and abundance reached 17.9 macroinvertebrates/kg. Significantly increased fine sediment deposition was detected within 1 km downstream of the restoration site and may negatively affect these habitats. Trade-offs between positive effects at restored sites and negative effects in downstream habitats need to be considered for a comprehensive evaluation of stream substratum restoration.     

Timing matters: species‐specific interactions between spawning time,substrate quality,and recruitment success in three salmonid species

Substratum quality and oxygen supply to the interstitial zone are crucial for the reproductive success of salmonid fishes. At present, degradation of spawning grounds due to fine sediment deposition and colmation are recognized as main factors for reproductive failure. In addition, changes in water temperatures due to climate change, damming, and cooling water inlets are predicted to reduce hatching success. We tested the hypothesis that the biological effects of habitat degradation depend strongly on the species‐specific spawning seasons and life‐history strategies (e.g., fall‐ vs. spring‐spawners, migratory vs. resident species) and assessed temperature as an important species‐specific factor for hatching success within river substratum. We studied the species‐specific differences in their responses to such disturbances using egg‐to‐fry survival of Danube Salmon (Hucho hucho), resident brown trout (Salmo trutta fario), and migratory brown trout (Salmo trutta lacustris) as biological endpoint. The egg incubation and hatching success of the salmonids and their dependence on temperature and stream substratum quality were compared. Hatching rates of Danube salmon were lower than of brown trout, probably due to higher oxygen demands and increased interstitial respiration in spring. Increases in maximum water temperature reduced hatching rates of resident and migratory brown trout (both fall‐spawners) but were positively correlated with hatching rates of Danube salmon (a spring‐spawner). Significantly longer incubation periods of resident and migratory brown trout coincided with relatively low stream substratum quality at the end of the egg incubation. Danube salmon seem to avoid low oxygen concentrations in the hyporheic zone by faster egg development favored by higher water temperatures. Consequently, the prediction of effects of temperature changes and altered stream substratum properties on gravel‐spawning fishes and biological communities should consider the observed species‐specific variances in life‐history strategies to increase conservation success.     

Species traits in relation to temporal and spatial heterogeneity in streams: a test of habitat templet theory

1. The habitat templet approach depends on defining templet axes appropriate to the organism(s) of interest, predicting the traits of species associated with different parts of the templet, and testing these predictions in a range of habitats whose positions in the templet have been determined. 2. In this study of thirty-five benthic insect taxa at fifty-four tributary sites of the Taieri River on the South Island of New Zealand, we chose as the temporal axis the intensity/frequency of disturbance, defined in terms of bed movement during high discharge events. As the spatial axis, we postulated that three features would provide refugia and therefore ameliorate disturbance—percentage of the bed with low shear stress, percentage of the bed made up of large substratum particles and availability of interstitial space in the bed—from which we derived a combined multivariate refugium axis. 3. More disturbed communities contained a significantly higher percentage of individuals possessing the following traits: small size, high adult mobility, habitat generalist (each predicted to confer resilience in response to disturbance), clinger, streamlined/flattened and with two or more life stages outside the stream (each predicted to confer resistance in the face of disturbance). When analyses were performed on the percentage of taxa having particular traits, the predicted positive relationships with average bed movement were found for high adult mobility and habitat generalist traits. 4. The percentage of variance in trait scores explained by intensity of disturbance was generally higher in sites with less refugia available and lower in sites further from the headwaters. The percentage of variance explained was higher in sites recently subject to a major high discharge disturbance, suggesting that disturbances tend to strengthen the pattern of preponderance of resilience/resistance traits. 5. We mapped insect taxa onto the two-dimensional templet, following Grime et al.’s triangular terrestrial plant classification. The full variety of resistance and resilience traits were represented in insect species throughout the templet, but taxa associated with more disturbed conditions generally displayed a larger number of resilience and resistance traits, combined, than taxa associated with more stable stream beds.     

Does Channel Incision Affect In‐stream Habitat? Examining the Effects of Multiple Geomorphic Variables on Fish Habitat

Incised river channels are dynamic components of fluvial systems, represent geomorphic degradation, and are encountered worldwide. Ecological effects of incision can be far‐reaching, affecting habitat availability and channel processes. Although incision can reflect habitat degradation, some studies suggest that important in‐stream habitats do not differ with the degree of incision. Therefore, we tested whether in‐stream habitat variables that are important to imperiled fishes differ in river reaches with varying degrees of incision. Because incision (measured using entrenchment ratio) had no discernable effect on in‐stream habitat characteristics (i.e., proportion fines, gravel, cobble, and macrophyte occurrence and length), we expanded our analysis to assess the effects of 29 additional geomorphic variables on in‐stream habitat. These analyses indicated that bank height, bed mobility, D₈₄, cross‐sectional area, bankfull width, and wetted perimeter accounted for 42% of macrophyte occurrence and 64% of macrophyte length variance. Postflood surveys indicated that macrophyte occurrence on cobble declined as bank height and bed mobility increased, and sediment size decreased, suggesting that sediment size and bed mobility have a stronger influence on in‐stream habitat than incision. Although channel incision often indicates environmental degradation, important aspects of habitat are not described by this measurement. Strategies that depend on incision to identify restoration sites may have limited habitat benefits in Southeastern Piedmont streams and rivers. Instead, landscape or shoal‐scale restoration approaches that increase coarse sediment proportions may increase macrophyte occurrence, length, and persistence. Sediment budgets that identify coarse and fine sediment sources and transport may be useful to prioritize restoration approaches.     

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.     

Egg-to-fry survival of the sea trout in some streams of Gotland

The quality of the surface water of Gotland (Sweden) streams was good, and did not limit sea trout egg-to-fry (ETF) survival. The oxygen concentration of the interstitial water was positively correlated to the permeability of the stream bed, and to the geometric mean diameter of the substratum. When the oxygen at 15 cm inside the stream bed was undiminished from surface values, the permeability of the stream bed was at least 6000 cm h^–1 and the geometric mean diameter of the substratum at least 15.0 mm. When the interstitial oxygen concentration increased, the interstitial ammonium concentration decreased, the NO₂ concentration remained stable and the NO₃ concentration increased. The ETF survival up to hatching was highly dependent upon the oxygen concentration of the interstitial water. For survival from hatching to emergence, the geometric mean diameter of the substratum was the most important factor. ETF survival >50% was observed where the interstitial oxygen concentration averaged at least 10mg I-¹, the substratum permeability at least 2000cm h-¹ and the substratum geometric mean diameter at least 15 mm. Other factors, such as overdigging and drying out of the spawning grounds could increase the ETF mortality.     

Habitat persistence for sedentary organisms in managed rivers: the case for the federally endangered dwarf wedgemussel (Alasmidonta heterodon) in the Delaware River

1. To manage the environmental flow requirements of sedentary taxa, such as mussels and aquatic insects with fixed retreats, we need a measure of habitat availability over a variety of flows (i.e. a measure of persistent habitat). Habitat suitability measures in current environmental flow assessments are measured on a ‘flow by flow’ basis and thus are not appropriate for these taxa. Here, we present a novel measure of persistent habitat suitability for the dwarf wedgemussel (Alasmidonta heterodon), listed as federally endangered in the U.S.A., in three reaches of the Delaware River. 2. We used a two‐dimensional hydrodynamic model to quantify suitable habitat over a range of flows based on modelled depth, velocity, Froude number, shear velocity and shear stress at three scales (individual mussel, mussel bed and reach). Baseline potentially persistent habitat was quantified as the sum of pixels that met all thresholds identified for these variables for flows ≥40 m³ s^?1, and we calculated the loss of persistently suitable habitat by sequentially summing suitable habitat estimates at lower flows. We estimated the proportion of mussel beds exposed at each flow and the amount of change in the size of the mussel bed for one reach. 3. For two reaches, mussel beds occupied areas with lower velocity, shear velocity, shear stress and Froude number than the reach average at all flows. In the third reach, this was true only at higher flows. Together, these results indicate that beds were possible refuge areas from the effects of these hydrological parameters. Two reaches showed an increase in the amount of exposed mussel beds with decreasing flow. 4. Baseline potentially persistent habitat was less than half the areal extent of potentially suitable habitat, and it decreased with decreasing flow. Actually identified beds and modelled persistent habitat showed good spatial overlap, but identified beds occupied only a portion of the total modelled persistent habitat, indicating either that additional suitable habitat is available or the need to improve habitat criteria. At one site, persistent beds (beds where mussels were routinely collected) were located at sites with stable substratum, whereas marginal beds (beds where mussels were infrequently collected or that were lost following a large flood event) were located in scoured areas. 5. Taken together, these model results support a multifaceted approach, which incorporates the effects of low and high flow stressors, to quantify habitat suitability for mussels and other sedentary taxa. Models of persistent habitat can provide a more holistic environmental flow assessment of rivers.     

Faunal structures associated with patches of mussels on East Asian coasts

Aggregations of mussels harbor a variety of associated animals and make it possible for diverse species to coexist at the shore. Species composition and diversity of the associated fauna are controlled by the position of mussel beds or patches, e.g. tidal level, age structure of mussels, quality of ambient water and by mussel species. When patches of mussels were surrounded by algal growth, a difference in the species composition of the associated fauna was recognized between the patches and algal mats. Mechanisms promoting coexistence are discussed. Biodeposit production by mussels may affect the environment both within the bed and the ambient waters. Reducing sediments showing low Eh values caused by the accumulation of biodeposits was observed in calm waters where the polychaete Capitella capitata, an indicator for organic enrichment, occurred both in the intertidal mussel bed and the subtidal sandy bottom communities. In a shallow subtidal sandy bottom of the Gulf of Thailand, where heavy bioturbation by the spatangoid urchin Brissus latecarinatus was occurring, small patches of the mussel Modiolus metcalfi increased species diversity and equitability in this habitat. Species composition was different between mussel patches and pure sandy bottoms. Electronic Publication     

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.     

Biotic resistance to invasion along an estuarine gradient

Biotic resistance is the ability of native communities to repel the establishment of invasive species. Predation by native species may confer biotic resistance to communities, but the environmental context under which this form of biotic resistance occurs is not well understood. We evaluated several factors that influence the distribution of invasive Asian mussels (Musculista senhousia) in Mission Bay, a southern California estuary containing an extensive eelgrass (Zostera marina) habitat. Asian mussels exhibit a distinct spatial pattern of invasion, with extremely high densities towards the back of Mission Bay (up to 4,000 m⁻²) in contrast with near-complete absence at sites towards the front of the bay. We established that recruits arrived at sites where adult mussels were absent and found that dense eelgrass does not appear to preclude Asian mussel growth and survival. Mussel survival and growth were high in predator-exclusion plots throughout the bay, but mussel survival was low in the front of the bay when plots were open to predators. Additional experiments revealed that consumption by spiny lobsters (Panulirus interruptus) and a gastropod (Pteropurpura festiva) likely are the primary factors responsible for resistance to Asian mussel invasion. However, biotic resistance was dependent on location within the estuary (for both species) and also on the availability of a hard substratum (for P. festiva). Our findings indicate that biotic resistance in the form of predation may be conferred by higher order predators, but that the strength of resistance may strongly vary across estuarine gradients and depend on the nature of the locally available habitat.     

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).     

The impact of land use on the mussel Margaritifera margaritifera and its host fish Salmo trutta

Today, land use impacts a major proportion of all streams. Here, landscape features in corridors along streams and water chemical factors were analyzed in relation to recruitment of the threatened freshwater pearl mussel (Margaritifera margaritifera) and its host fish the brown trout (Salmo trutta). Mussel recruitment and trout density were negatively related to forest clear-cuts. Mussel recruitment was negatively related to water color and turbidity. Therefore, the threats to the mussel may be severe, as low mussel recruitment may be caused by direct effects on the juvenile mussels and indirect effects on the host fish. High proportions of lakes and ponds were found to be positive for recruitment and for trout, and deciduous forest was positively related to trout. The combination of investigations at different scales at the landscape level and at in-stream levels may be applicable to find threats to other threatened species. The results indicate that forestry activities may negatively affect recruitment of freshwater pearl mussels and its host fish. Reductions of forestry activities and the retaining of intact quantity and quality of the riparian zones next to streams, both for the mussel and its host fish may be important conservation measures to restore freshwater pearl mussel populations.     

Influence of Bed‐Sediment Features on the Interstitial Habitat Available for Macroinvertebrates in 15 French Streams

The interstitial space of bed‐sediments is a major component of the stream system as it influences both physical and biological processes and patterns, such as exchanges between surface water and groundwater or the assemblage of invertebrates. To investigate the influence of grain‐size distribution and vertical bed‐packing on the interstitial habitat, we used 99 frozen cores collected from 15 French streams. On our data, overall invertebrate density, density of Chironomidae and taxon richness were more strongly related to effective porosity than to other physical variables describing bed‐sediments. We built two models to quantify the influence of grain‐size distribution and vertical bed‐packing on effective porosity. The effective porosity decreased with increasing grain‐size heterogeneity and was strongly related to grain‐size distribution. These relationships differed with vertical bed‐packing, but were consistent across three geologic types of sediments that varied in particle shape and particle texture. These general relationships suggest that effective porosity could be helpful to better understand the distribution patterns of invertebrates in bed‐sediments.     

Gravel dredging alters diversity and structure of riverine fish assemblages

1. Human activities affect fish assemblages in a variety of ways. Large‐scale and long‐term disturbances such as in‐stream dredging and mining alter habitat and hydrodynamic characteristics within rivers which can, in turn, alter fish distribution. Habitat heterogeneity is decreased as the natural riffle–pool–run sequences are lost to continuous pools and, as a consequence, lotic species are displaced by lentic species, while generalist and invasive species displace native habitat specialists. Sediment and organic detritus accumulate in deep, dredged reaches and behind dams, disrupting nutrient flow and destroying critical habitat for habitat specialist species. 2. We used standard ecological metrics such as species richness and diversity, as well as stable isotope analysis of δ¹³C and δ¹⁵N, to quantify the differences in fish assemblages sampled by benthic trawls among dredged and undredged sites in the Allegheny River, Pennsylvania, U.S.A. 3. Using mixed‐effects models, we found that total catch, species richness and diversity were negatively correlated with depth (P < 0.05), while species richness, diversity and proportion of species in lithophilic (‘rock‐loving’) reproductive guilds were lower at dredged than at undredged sites (P < 0.05). 4. Principal components analysis and manova revealed that taxa such as darters in brood hider and substratum chooser reproductive guilds were predominantly associated with undredged sites along principal component axis 1 (PC1 and manova P < 0.05), while nest spawners such as catfish and open substratum spawners including suckers were more associated with dredged sites along PC2 (P < 0.05). 5. Stable isotope analysis of δ¹³C and δ¹⁵N revealed shifts from reliance on shallow water and benthic‐derived nutrients at undredged sites to reliance on phytoplankton and terrestrial detritus at deep‐water dredged sites. Relative trophic positions were also lower at dredged sites for many species; loss of benthic nutrient pathways associated with depth and dredging history is hypothesised. 6. The combination of ecological metrics and stable isotope analysis thus shows how anthropogenic habitat loss caused by gravel dredging can decrease benthic fish abundance and diversity, and that species in substratum‐specific reproductive guilds are at particular risk. The effects of dredging also manifest by altering resource use and nutrient pathways within food webs. Management and conservation decisions should therefore consider the protection of relatively shallow areas with suitable substratum for spawning for the protection of native fishes.     

Density‐dependent mortality in adults,but not juveniles,of stream‐resident brown trout (Salmo trutta)

1. This study investigates when and where density dependence operates on the mortality rates of stream‐resident brown trout Salmo trutta. To this aim, I explored populations in habitats of different quality containing high, low or intermediate densities over broad scales of space and time. The study is based on census data of 170 cohorts quantified from recruitment to the total disappearance at 12 sites in four contrasting tributaries of the Rio Esva drainage (north‐western Spain), over the years 1986–2007. 2. Log₁₀‐transformed survivor density over time highlighted a consistent pattern for the 170 cohorts characterised by the occurrence of only two life stages. An early stage starts at recruitment, lasts about half the lifetime and shows no or negligible mortality. A threshold time at 425–620 days after emergence preceded a second stage of continuous and constant mortality until the final disappearance of the cohorts. Consequently, in all scenarios, mortality only occurred in the adult component and no effect of season, year, age‐class and/or reproductive stage was detected. 3. Substantial spatial and temporal variations typified both recruitment (range R = 0.01–1.62 ind m^?2) and adults’ mortality rates (range Z = 0.03–0.38 day^?1). Nested anova s revealed strong effects of site and year on both recruitment and mortality with sites interspersed along the stream gradients where recruitment and mortality were typically high relative to other sites located either nearby in the same stream or distant in another stream, where both recruitment and mortality rates were typically low or intermediate. 4. Adult mortality rates plotted against recruitment for the 170 cohorts pooled revealed a continuous, positive power relationship that explained 45.3% of variation in mortality rates over the whole range of recruitment values. Similarly, highly significant power relationships were elucidated for site‐specific mortality rates averaged across years and for annual‐specific mortality rates averaged across sites against the corresponding mean recruitment averaged across years and sites, respectively. These relationships support the hypothesis that the operation of density dependence is scale independent and context independent but operates in a continuous manner across all scenarios examined. 5. A chronic effect of density dependence on adult losses induces temporally persistent populations maintained by a low number of spawners. Apparently, the operation of density dependence adjusts the number of spawners to the availability of rearing and spawning habitat. This dynamic process may also help to explain the small effective population size (Ne) recently documented by genetic studies of stream‐living brown trout and other salmonids.