Hydrologic Connectivity Affects Fish Assemblage Structure,Diversity, and Ecological Traits in the Unregulated Gambia River,West Africa

The Gambia River of West Africa is a large unobstructed river, characterized by a natural flow regime and lateral connectivity across its floodplain. Construction of a major dam, however, is planned. We compared patterns of fish diversity, habitat use, assemblage structure, and the distribution of trophic position and body morphology in riverine and floodplain habitats in Niokolo Koba National Park, located downstream of the planned dam site. A total of 49 fish species were captured, revealing a lognormal distribution as expected for species‐rich assemblages. Fish species exhibited a range of habitat use patterns, from generalist to highly habitat‐specific, and appeared to migrate laterally among habitats between seasons. Species richness was homogenous among habitats in the wet season yet appeared to increase with isolation from the main river in the dry season. Fish assemblage structure was best explained by the interaction between habitat type and season, underlining the importance of the natural flow regime and lateral connectivity among floodplain habitats. The abundance of fishes having elongate bodies increased with isolation from the main channel in the wet season only. The distribution of fishes having compressed cross‐sectional morphology decreased with isolation from the main channel in the dry season only. These patterns of trait distribution support the conclusion that variation in hydrologic connectivity structures the fish assemblage. Our results suggest that altered flow regimes and loss of floodplain habitats after damming could lead to both decreased taxonomic and functional diversity of the fish assemblage.     

The contribution of long‐term isolated water bodies to floodplain fish diversity

1. Floodplains and their water bodies are typical ecotopes of large lowland rivers. The lowland Oder River, Germany, provided a rare opportunity to study fish assemblages of comparable floodplain water bodies differing by >50 years of isolation history. We hypothesised that true floodplain specialists peak in rarely connected water bodies, while frequently flooded waters support tolerant generalists. 2. Three macrohabitats, main channel (MC), temporarily connected and isolated floodplain water bodies, were sampled by electro‐fishing and their fish assemblage characters recorded. 3. Long‐term isolation of floodplain water bodies had a significant effect on the fish assemblage by promoting species preferring still water. Limnophilic and floodplain specialist species significantly increased with isolation. 4. Fish densities, species richness and diversity clearly differed between MC sites and floodplain water bodies. Shannon’s species diversity index peaked in both the MC and isolated waters and was lowest in the temporarily flooded waters where eurytopic fish dominated. 5. The significant gain in abundance and numbers of limnophilic species in the isolated compared to the temporarily flooded water bodies underpinned the significant contribution of long‐term isolated waters to the gamma‐diversity of large floodplain rivers, which should be considered in floodplain rehabilitation.     

Comparative use of side and main channels by small‐bodied fish in a large,unimpounded river

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Geomorphological characteristics of the Wabash River,USA: Influence on fish assemblages

River hydrogeomorphology is a potential predictor of ecosystem and assemblage variation. We tested for fish assemblage variation as a function of hydrogeomorphology in a Midwestern US large river, the Wabash River. Fish data were classified by taxonomy and traits and we tested if assemblages varied with river hydrogeomorphology or river distance, defined into 10‐km distinct reaches. Three unique geomorphological units, Functional Process Zones (FPZ), were identified using an ArcGIS hydrogeomorphic model, based primarily on channel width, floodplain width, and down valley slope. Five locations were identified as FPZ A with narrow stream channel, high down valley slope, and an expansive floodplain. Ten locations were identified as FPZ B with a wide river channel and wide floodplain. Thirty‐five locations were identified as FPZ C with wide river channel and a constrained floodplain. The sites were categorized into three stream orders: 5, 6, and 7. We found hydrogeomorphology classified by unique FPZs or by river distance influenced taxonomic and functional fish assemblages for the Wabash River. There was high overlap among fish occurrences among FPZs, but nine species resulted as significant indicators of specific FPZs. Five traits were significant indicators of FPZs: an intermediate Swim Factor score, medium tolerance to silt, small‐large stream size preference, and two Shape Factor categories. Our conclusions are that fish assemblages respond strongly to local geomorphology and river distance, fitting the riverine ecosystem synthesis and the river continuum concept.     

Diversity,occurrence and feeding traits of caddisfly larvae as indicators for ecological integrity of river-floodplain habitats along a connectivity gradient

In order to assess ecological values of Lower Rhine and Meuse floodplain habitats we studied the spatial and seasonal variation in diversity, species assemblages and feeding traits of caddisfly larvae in water bodies over the lateral connectivity gradient: eupotamon: main and secondary channels; parapotamon: channels connected permanently with the main channel only at their downstream ends; plesiopotamon: disconnected channels close to the main channel; paleopotamon: abandoned meanders at a greater distance from the main channel.Spatial variety was studied by analyzing the summer species composition in 70 Lower Rhine and Meuse water bodies which were categorized in connectivity habitats, whereas seasonal variety was studied in Lower Rhine water bodies along a connectivity gradient by monthly sampling over a whole year. Physico-chemical data and environmental parameters were recorded for each water body during sampling. Diversity and species assemblages of caddisfly larvae varied in relation to connectivity, macrophyte diversity and abundance and stream velocity. A comparison with historical records and species lists from less disturbed rivers showed that diversity in the main channel was very low.Caddisfly larvae species assemblages varied over the connectivity gradient. Lotic habitats (eupotamon) were separated from the lentic ones, and the well vegetated paleopotamon from the sparsely vegetated parapotamon and pleisopotamon habitats, indicating the overall importance of vegetation and current velocity for the species assemblages. Hydropsychidae have been found in the eupotamon exclusively, whereas Limnephilidae, Hydroptilidae and Polycentropodidae have been found predominantly in the paleopotamon water bodies. Leptoceridae were found in all floodplain water body categories. A similar pattern of distribution of families along the lateral connectivity gradient was found in more natural rivers.Caddisfly larvae species feeding traits showed a clear differentiation over the lateral connectivity gradient with filter-feeders and scrapers most important in the eupotamon and parapotamon, and shredders, piercers and predators most dominant in the paleopotamon habitats, indicating the importance of nutritional resources in relation to hydrological connectivity for the structure and functioning of caddisfly larvae species assemblages. The analysis of the species feeding traits allows generalizations towards the entire aquatic community and general prognoses for other floodplain ecosystems.     

A multi-faceted framework of diversity for prioritizing the conservation of fish assemblages

Floodplain waterbodies and their biodiversity are increasingly threatened by human activities. Given the limited resources available to protect them, methods to identify the most valuable areas for biodiversity conservation are urgently needed. In this study, we used freshwater fish assemblages in floodplain waterbodies to propose an innovative method for selecting priority areas based on four aspects of their diversity: taxonomic (i.e. according to species classification), functional (i.e. relationship between species and ecosystem processes), natural heritage (i.e. species threat level), and socio-economic (i.e. species interest to anglers and fishermen) diversity. To quantitatively evaluate those aspects, we selected nine indices derived either from metrics computed at the species level and then combined for each assemblage (species rarity, origin, biodiversity conservation concern, functional uniqueness, functional originality, fishing interest), or from metrics directly computed at the assemblage level (species richness, assemblage rarity, diversity of biological traits). Each of these indices belongs to one of the four aspects of diversity. A synthetic index defined as the sum of the standardized aspects of diversity was used to assess the multi-faceted diversity of fish assemblages. We also investigated whether the two main environmental gradients at the catchment (distance from the sea) and at the floodplain (lateral connectivity of the waterbodies) scales influenced the diversity of fish assemblages, and consequently their potential conservation value. Finally, we propose that the floodplain waterbodies that should be conserved as a priority are those located in the downstream part of the catchment and which have a substantial lateral connectivity with the main channel.     

The Significance of Meander Restoration for the Hydrogeomorphology and Recovery of Wetland Organisms in the Kushiro River,a Lowland River in Japan

The meanders and floodplains of the Kushiro River were restored in March 2011. A 1.6‐km stretch of the straightened main channel was remeandered by reconnecting the cutoff former channel and backfilling the straightened reach, and a 2.4‐km meander channel was restored. Additionally, flood levees were removed to promote river–floodplain interactions. There were four objectives of this restoration project: to restore the in‐stream habitat for native fish and invertebrates; to restore floodplain vegetation by increasing flooding frequency and raising the groundwater table; to reduce sediment and nutrient loads in the core wetland areas; to restore a river–floodplain landscape typical to naturally meandering rivers. In this project, not only the natural landscape of a meandering river but also its function was successfully restored. The monitoring results indicated that these goals were likely achieved in the short term after the restoration. The abundance and species richness of fish and invertebrate species increased, most likely because the lentic species that formerly inhabited the cutoff channel remained in the backwater and deep pools created in the restored reach. In addition, lotic species immigrated from neighboring reaches. The removal of flood levees and backfilling of the formerly straightened reach were very effective in increasing the frequency of flooding over the floodplains and raising the water table. The wetland vegetation recovered rapidly 1 year after the completion of the meander restoration. Sediment‐laden floodwater spread over the floodplain, and approximately 80–90% of the fine sediment carried by the water was filtered out by the wetland vegetation.     

Influence of stream geomorphic condition on fish communities in Vermont, U.S.A.

1. Evaluations of stream geomorphic condition may increase our understanding of the composite effects of human‐induced habitat change on fish communities. Using systematic sampling of 44 reaches spread across 26 rivers in Vermont from 2002 through 2004, we tested the hypothesis that stream reaches in reference geomorphic condition would support fish assemblages that differed in diversity and productivity from fish communities found in reaches of poorer geomorphic condition. 2. At each study reach, we sampled the fish community, identified the morphological unit according to common stream classification systems and then evaluated the extent of deviation from reference geomorphic condition using a regionally adapted geomorphic assessment methodology. 3. We used principal component analysis (PCA) and linear regression to build exploratory models linking stream geomorphic condition to fish community characteristics. 4. Our results suggest that geomorphic condition significantly influences fish community diversity, productivity and condition. Geomorphic condition was a significant factor in all of our fish community models. In conjunction with additional reach characteristics, geomorphic condition explained up to 31% of the total variance observed in models for species diversity of fish communities, 44% of the variance in assemblage biomass and 45% of the variance in a regional index of biotic integrity. 5. Our work builds on single‐species evidence that geomorphic characteristics represent important local‐scale fish‐habitat variables, showing that stream geomorphic condition is a dominant factor affecting entire fish communities. Our results enhance our understanding of the hierarchy of factors that influences fish community diversity and organisation and support the use of geomorphic condition assessments in stream management.     

Estimation of minimum area requirement of river‐connected lakes for fish diversity conservation in the Yangtze River floodplain

Aim Hydrological disconnection of floodplains from rivers is among the top factors threatening river‐floodplain ecosystems. To keep enough floodplain area is of great importance to biodiversity conservation. In the Yangtze River floodplain, most lakes were disconnected from the mainstream by dams in 1950–1970s. By analysing fish diversity data, we aim at determining the effects of river‐lake disconnection on fish diversity, at estimating the minimum protected area of river‐connected lakes and at proposing a holistic strategy for fish conservation in the mid‐lower reaches of the river. Location The Yangtze River floodplain, China. Methods We collected recorded data of fish diversity of 30 Yangtze floodplain lakes. Species–area relationships were analysed and compared between river‐connected and river‐disconnected lakes. Cumulative species–area models were constructed to estimate the minimum protected area of river‐connected lakes. Results River‐lake disconnection reduced fish diversity of Yangtze lakes by 38.1%, so that the river‐connected lakes play an important role in maintaining the floodplain biodiversity. The minimum protected area of river‐connected lakes was estimated to be 14,400 km². Therefore, we should not only protect the existent connected lakes of 5500 km², but also reconnect disconnected lakes of at least 8900 km² in the Yangtze basin. Main conclusions Species–area relationships are of importance in reserve design. We suggest that cumulative species–area model might be more suitable for ecosystems with high connectivity among regions such as floodplains. As the Yangtze River floodplain is an integrative ecosystem, we suggest establishing a holistic nature reserve in the mid‐lower basin for effective conservation of biodiversity.     

Analysis of the reproductive strategy of <Emphasis Type="Italic">Jenynsia multidentata</Emphasis> (Cyprinodontiformes,Anablepidae) with focus on sexual differences in growth,size, and abundance

The habitat heterogeneity generated and sustained by the connectivity of floodplain habitats, the seasonal flood pulse, and the variability of the physical structures typically found in floodplains of large rivers results in a variable space–time mosaic of water sources that results in a high biodiversity of the river-floodplain system. In order to assess the implications of natural connectivity and the heterogeneity on the patterns of macroinvertebrate assemblages at different spatial scales, monthly samplings in six different mesohabitats (lakes with different hydrological connection and secondary channels with permanently and intermittent flow) of the Paraná River floodplain were performed from April 2005 to March 2006. The mesohabitats had different granulometry and detritus composition of their bottom sediments. They also had different conductivity, transparency, and depth in relation to the different connectivity degrees. Mesohabitats differed in the abundance of macroinvertebrates of different taxonomic groups and diversity. The environmental variables were correlated to the patterns of macroinvertebrate abundance, with dominance of different species of annelids and mollusks at the patch, mesohabitats, and island scales. An alpha diversity gradient from the isolated lake (65 taxonomic units) to the secondary channels (25 taxonomic units) was obtained. The analyzed mesohabitats showed a high taxa turnover, with high values not only among the mesohabitats located in the different islands, but also among the mesohabitats in relation to different connectivity degrees. The mesohabitats showed negative co-occurrence of macroinvertebrate assemblages. The spatial heterogeneity, sustained by the connectivity degree, played a key role in structuring benthic assemblages at different scales, positively influencing the regional diversity.     

Relationships between fish assemblages, macrophytes and environmental gradients in the Amazon River floodplain

During the flood season of 1992–1993, 139 species of fishes were collected from a floodplain lake system in the central Amazon Basin. Fish species distribution was examined relative to abiotic variables in seven vegetation strata on Marchantaria Island, Solimões River. Both environmental variables and species distributions were influenced by a river channel to floodplain-interior gradient. Species diversity was significantly higher in vegetated areas than in unvegetated areas, with deeper water Paspalum repens stands harbouring the highest diversity. As a result, species richness and catches were positively related to habitat complexity, while catch was also negatively related to dissolved oxygen (DO) and water depth. Low DO and shallow waters appeared to act as a refuge from predation. Fish assemblages were related to water chemistry, but species richness was not. Canonical correspondence analysis provided evidence that floodplain fish assemblages formed by the 76 most common species were influenced by physical variables, macrophyte coverage and habitat complexity, which jointly accounted for 67% of the variance of fish species assemblages. Omnivores showed no pattern relative to the river channel to floodplain-interior gradient while detritivores were more likely to be found at interior floodplain sites and piscivores closer to the river. Piscivores could be further separated into three groups, one with seven species associated with free-floating macrophytes in deep water, a second with five species found in shallow waters with rooted grasses and a third with six open water orientated species. The results suggest that fish assemblages in the Amazon floodplain are not random associations of species.     

Altitudinal gradients in stream fish diversity and the prevalence of diadromy in the Sixaola River basin,Costa Rica

Landscape-scale patterns of freshwater fish diversity and assemblage structure remain poorly documented in many areas of Central America, while aquatic ecosystems throughout the region have been impacted by habitat degradation and hydrologic alterations. Diadromous fishes may be especially vulnerable to these changes, but there is currently very little information available regarding their distribution and abundance in Central American river systems. We sampled small streams at 20 sites in the Sixaola River basin in southeastern Costa Rica to examine altitudinal variation in the diversity and species composition of stream fish assemblages, with a particular focus on diadromous species. A set of environmental variables was also measured in the study sites to evaluate how changes in fish assemblage structure were related to gradients in stream habitat. Overall, fish diversity and abundance declined steeply with increasing elevation, with very limited species turnover. The contribution of diadromous fishes to local species richness and abundance increased significantly with elevation, and diadromous species dominated assemblages at the highest elevation sites. Ordination of the sampling sites based on fish species composition generally arranged sites by elevation, but also showed some clustering based on geographic proximity. The dominant gradient in fish community structure was strongly correlated with an altitudinal habitat gradient identified through ordination of the environmental variables. The variation we observed in stream fish assemblages over relatively small spatial scales has significant conservation implications and highlights the ecological importance of longitudinal connectivity in Central American river systems.     

Local and ecoregion effects on fish assemblage structure in tributaries of the Rio Paraíba do Sul, Brazil

1.  We examined the effects of physical and chemical habitat variables and ecoregions on species occurrence and fish assemblage structure in streams of the Paraíba do Sul basin, in southeast Brazil.
2.  Fish and environmental data were collected from 42 sites on 26 first to fourth order streams (1 : 50 000 map scale) in three ecoregions. The sites occurred in one valley and two plateau ecoregions at altitudes of 40–1080 m and distances of 0.1–188 km from the main channel of the Rio Paraíba do Sul. Physical habitat (substratum, riparian cover, habitat types) and water quality (dissolved oxygen, pH, temperature and conductivity) variables were measured at each sampling site.
3.  A total of 2684 individuals in 16 families and 59 species were recorded.
4.  Ecoregion was a better predictor of the fish assemblage than the other environmental variables, according to the differences between the mean within-class and mean between-class similarities in assemblage data.
5.  Differing landscape characteristics were associated with differing local variables and thereby with differing fish assemblage structures. Riffles, shrub, grass, dissolved oxygen, conductivity and temperature were closely related to fish assemblage structure.
6.  Fish assemblages in sites far from the main river and at higher altitudes also differed from those near the Paraíba do Sul main channel, presumably as a result of differences in connectivity, covarying environmental factors and anthropogenic influence.
7.  These results reinforce the importance of understanding how stream communities are influenced by processes and patterns operating at local and regional scales, which will aid water resource managers to target those factors in their management and rehabilitation efforts.     

Patterns of fish species richness in the Seine River basin,France

Variation in fish assemblage structure was examined in different rivers of the Seine Basin. Factor analysis was used to identify similarities among samples and species, and to show ichtyological changes along an upstream-downstream gradient. Fish species richness was correlated with catchment area, distance from the source and number of individuals. Relationships between species richness and number of individuals in a sample were largely an artifact of sampling.Species richness increased with river size, reached a maximum in midsize rivers, then decreased in large rivers. This species richness pattern was consistent with the model of the River Continuum Concept. However, possible causes could include greater pollution effects in large rivers. Distribution of feeding guilds was related to river size. Species richness and proportion of omnivores and piscivores increased with river size, whereas species richness and proportion of invertivores declined downstream. These patterns suggest that trophic diversity of fish assemblages may be related to food availability.     

Spatial and temporal variation of fish assemblages and their associations to habitat variables in a mountain stream of north Tiaoxi River,China

The spatial and temporal variations of the fish assemblages in mountain streams of China are poorly understood. The relationships between the fish assemblage and selected habitat features were examined in the North Tiaoxi River, one of headwaters of Taihu Lake. A total of 3,348 individuals belonging to 5 orders, 11 families, 25 genera and 34 species were collected including 33 native species and one invasive species. Among those, about 20 species were endemic to China. Non-metric Multidimensional Scaling (NMDS) was applied to compare fish assemblage structures from upstream to downstream during four seasons. Species assemblages differed along the stream continuum, but there was little apparent change associated with the seasons. Species richness and Shannon-Weaver index (H′) tended to increase along the stream continuum from the upstream to downstream and the proportion of invertivorous fish tended to significantly decrease along the continuum with a parallel significant increase in the percentage of omnivores. Fish assemblages were significantly related to both water quality and habitat structure variables. Canonical Correspondence Analysis ordinations (CCA) revealed that 6 of the 14 selected environmental variables had significant relationships with the fish assemblage such as distance to source, stream width, altitude, pH, water depth, and water velocity and different sampling sites were associated with different environmental variables in different seasons. The main differences in fish assemblage structure and diversity within the whole watercourse are probably related to large-scale factors such distance to source, altitude and stream width. Differences of instream characteristics are likely to be caused by natural variability of the ecosystems but also, in some case, by anthropogenic influence like human settlements, agriculture and river embankment and pollution from small factory.     

Influences of low-head dams on the fish assemblages in the headwater streams of the Qingyi watershed, China

The influences of low-head dams on the fish assemblages were examined in this study, using fish data collected in six treatment and five reference sites at three low-head dams in the headwater streams of the Qingyi watershed, China. Comparing with those in the reference sites, local habitat variables were significantly altered by low-head dams in the treatment sites, involving wider channel (only in the impoundment area), deeper water and slower flow. Fish species richness varied significantly across seasons, not across site categories, suggesting that these low-head dams did not alter species richness. However, significant decreases in fish abundance and density were observed in the impoundment areas immediately upstream of dams, but not in the plunge areas downstream. Fish assemblage structures kept relative stability across seasons, and their significant difference between-site was only observed between the impoundment areas and the sites far from dams upstream. This variation in assemblage structures was due to the differing relative abundance of some co-occurring species; more lentic but less lotic fish was observed in the impoundment areas. The spatial and temporal patterns of fish assemblages were correlated with local habitat in this study area. Wetted width had negative correlation with fish species richness, abundance and density, respectively. Water temperature also positively affected species richness. In addition, wetted width, water depth, current velocity and substrate were the important habitat variables influencing assemblage structures. Our results suggested that, by modifying local habitat characteristics, low-head dams altered fish abundance and density in the impoundment areas immediately upstream of dam, not in the plunge areas immediately downstream, and thereby influenced fish assemblage structures in these stream segments.     

Linking Flow Regime,Floodplain Lake Connectivity and Fish Catch in a Large River-Floodplain System,the Volga–Akhtuba Floodplain (Russian Federation)

River-floodplain systems are amongst the most productive—but often severely impacted—aquatic systems worldwide. We explored the ecological response of fish to flow regime in a large river-floodplain system by studying the relationships between (1) discharge and inundated floodplain area, with a focus on spatial and temporal patterns in floodplain lake connectivity, and (2) flood volume and fisheries catch. Our results demonstrate a non-linear relationship between discharge and floodplain inundation with considerable hysteresis due to differences in inundation and drainage rate. Inundation extent was mostly determined by flood volume, not peak discharge. We found that the more isolated lakes (that is, lakes with a shorter connection duration to the river) are located at higher local elevation and at larger hydrological distance from the main rivers: geographical distance to the river appears a poor predictor of lake isolation. Although year-to-year fish catches in the floodplain were significantly larger with larger flood volumes in the floodplain, they were not in the main river, suggesting that mechanisms that increase catch, such as increased floodplain access or increased somatic growth, are stimulated by flooding in the floodplain, but not in the river. Fish species that profit from flooding belong to different feeding guilds, suggesting that all trophic levels may benefit from flooding. We found indications that the ecological functioning of floodplains is not limited to its temporary availability as habitat. Refugia can be present within the floodplain itself, which should be considered in the management of large rivers and their floodplain.     

Consistent trophic patterns among fishes in lagoon and channel habitats of a tropical floodplain river: Evidence from stable isotopes

The relationship between food web dynamics and hydrological connectivity in rivers should be strongly influenced by annual flood pulses that affect primary production dynamics and movement of organic matter and consumer taxa. We sampled basal production sources and fishes from connected lagoons and the main channel of a low-gradient, floodplain river within the Orinoco River Basin in Venezuela. Stable isotope analysis was used to model the contribution of four basal production sources to fishes, and to examine patterns of mean trophic position during the falling-water period of the annual flood cycle. IsoSource, a multi-source mixing model, indicated that proportional contributions from production sources to fish assemblages were similar in lagoons and the main channel. Although distributions differed, the means for trophic positions of fish assemblages as well as individual species were similar between the two habitats. These findings contradict recent food web studies conducted in temperate floodplain rivers that described significant differences in trophic positions of fishes from slackwater and floodplain versus main channel habitats. Low between-habitat trophic variation in this tropical river probably results from an extended annual flood pulse (ca. 5 mo.) that allows mixing of sestonic and allochthonous basal production sources and extensive lateral movements of fishes throughout the riverscape.     

Method-integrated fish assemblage structure at two spatial scales along a free-flowing stretch of the Austrian Danube

Fish assemblages in large rivers are highly diverse communities. However, a single method to describe these complex systems adequately is not available as of yet. In this paper we propose a combination of normalized catch-per-unit-effort data of three fish sampling methods, thus providing a more inclusive representation of the overall assemblage. We investigate fish species occurrence and abundance in a survey at three consecutive sites in the free-flowing main channel of the Austrian Danube east of Vienna. Using boat and wading electrofishing, as well as longline fishing we collected data in four mesohabitats at each site. We analysed richness, assemblage structure and abundance of single species and tested site and mesohabitat as differentiating forces on the fish assemblage structure, applying PERMANOVA and NMDS. The investigated sites are populated by a species-rich fish assemblage, of which we collected a total of 36 species. The composition of the assemblage was rather consistent except for the occasional occurrence of rare species. At the smaller mesohabitat scale, however, the assemblages differed considerably and hydromorphological conditions were apparently the major structuring force. Overall, the combination of methods considerably improves the assessment across sites and mesohabitats and contributes to our understanding of large river fish assemblages.