Quantifying relationships between land‐use gradients and structural and functional indicators of stream ecological integrity

1. Modification of natural landscapes and land‐use intensification are global phenomena that can result in a range of differing pressures on lotic ecosystems. We analysed national‐scale databases to quantify the relationship between three land uses (indigenous vegetation, urbanisation and agriculture) and indicators of stream ecological integrity. Boosted regression tree modelling was used to test the response of 14 indicators belonging to four groups – water quality (at 578 sites), benthic invertebrates (at 2666 sites), fish (at 6858 sites) and ecosystem processes (at 156 sites). Our aims were to characterise the ecological response curves of selected functional and structural metrics in relation to three land uses, examine the environmental moderators of these relationships and quantify the relative utility of metrics as indicators of stream ecological integrity. 2. The strongest indicators of land‐use effects were nitrate + nitrite, delta‐15 nitrogen value (δ¹⁵N) of primary consumers and the Macroinvertebrate Community Index (a biotic index of organic pollution), while the weakest overall indicators were gross primary productivity, benthic invertebrate richness and fish richness. All indicators declined in response to removal of indigenous vegetation and urbanisation, while variable responses to agricultural intensity were observed for some indicators. 3. The response curves for several indicators suggested distinct thresholds in response to urbanisation and agriculture, specifically at 10% impervious cover and at 0.1 g m^?3 nitrogen concentration, respectively. 4. Water quality and ecosystem process indicators were influenced by a combination of temperature, slope and flow variables, whereas for macroinvertebrate indicators, catchment rainfall, segment slope and temperature were significant environmental predictor variables. Downstream variables (e.g. distance to the coast) were significant in explaining residual variation in fish indicators, not surprisingly given the preponderance of diadromous fish species in New Zealand waterways. The inclusion of continuous environmental variables used to develop a stream typology improved model performance more than the inclusion of stream type alone. 5. Our results reaffirm the importance of accounting for underlying spatial variation in the environment when quantifying relationships between land use and the ecological integrity of streams. Of distinctive interest, however, were the contrasting and complementary responses of different indicators of stream integrity to land use, suggesting that multiple indicators are required to identify land‐use impact thresholds, develop environmental standards and assign ecological scores for reporting purposes.     

Disentangling the pathways of land use impacts on the functional structure of fish assemblages in Amazon streams

Agricultural land use is a primary driver of environmental impacts on streams. However, the causal processes that shape these impacts operate through multiple pathways and at several spatial scales. This complexity undermines the development of more effective management approaches, and illustrates the need for more in‐depth studies to assess the mechanisms that determine changes in stream biodiversity. Here we present results of the most comprehensive multi‐scale assessment of the biological condition of streams in the Amazon to date, examining functional responses of fish assemblages to land use. We sampled fish assemblages from two large human‐modified regions, and characterized stream conditions by physical habitat attributes and key landscape‐change variables, including density of road crossings (i.e. riverscape fragmentation), deforestation, and agricultural intensification. Fish species were functionally characterized using ecomorphological traits describing feeding, locomotion, and habitat preferences, and these traits were used to derive indices that quantitatively describe the functional structure of the assemblages. Using structural equation modeling, we disentangled multiple drivers operating at different spatial scales, identifying causal pathways that significantly affect stream condition and the structure of the fish assemblages. Deforestation at catchment and riparian network scales altered the channel morphology and the stream bottom structure, changing the functional identity of assemblages. Local deforestation reduced the functional evenness of assemblages (i.e. increased dominance of specific trait combinations) mediated by expansion of aquatic vegetation cover. Riverscape fragmentation reduced functional richness, evenness and divergence, suggesting a trend toward functional homogenization and a reduced range of ecological niches within assemblages following the loss of regional connectivity. These results underscore the often‐unrecognized importance of different land use changes, each of which can have marked effects on stream biodiversity. We draw on the relationships observed herein to suggest priorities for the improved management of stream systems in the multiple‐use landscapes that predominate in human‐modified tropical forests.     

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.     

Temporal trends in algae, benthic invertebrate, and fish assemblages in streams and rivers draining basins of varying land use in the south-central United States, 1993–2007

Site-specific temporal trends in algae, benthic invertebrate, and fish assemblages were investigated in 15 streams and rivers draining basins of varying land use in the south-central United States from 1993–2007. A multivariate approach was used to identify sites with statistically significant trends in aquatic assemblages which were then tested for correlations with assemblage metrics and abiotic environmental variables (climate, water quality, streamflow, and physical habitat). Significant temporal trends in one or more of the aquatic assemblages were identified at more than half (eight of 15) of the streams in the study. Assemblage metrics and abiotic environmental variables found to be significantly correlated with aquatic assemblages differed between land use categories. For example, algal assemblages at undeveloped sites were associated with physical habitat, while algal assemblages at more anthropogenically altered sites (agricultural and urban) were associated with nutrient and streamflow metrics. In urban stream sites results indicate that streamflow metrics may act as important controls on water quality conditions, as represented by aquatic assemblage metrics. The site-specific identification of biotic trends and abiotic–biotic relations presented here will provide valuable information that can inform interpretation of continued monitoring data and the design of future studies. In addition, the subsets of abiotic variables identified as potentially important drivers of change in aquatic assemblages provide policy makers and resource managers with information that will assist in the design and implementation of monitoring programs aimed at the protection of aquatic resources.     

Implications of community concordance for assessing stream integrity at three nested spatial scales in Minnesota,U.S.A.

1. Fish and invertebrate assemblage data collected from 670 stream sites in Minnesota (U.S.A.) were used to calculate concordance across three nested spatial scales (statewide, ecoregion and catchment). Predictive taxa richness models, calibrated using the same data, were used to evaluate whether concordant communities exhibited similar trends in human‐induced taxa loss across all three scales. Finally, we evaluated the strength of the relationship between selected environmental variables and the composition of both assemblages at all three spatial scales. 2. Significant concordance between fish and invertebrate communities occurred at the statewide scale as well as in six of seven ecoregions and 17 of the 21 major catchments. However, concordance was not consistently indicative of significant relationships between rates of fish and invertebrate taxa loss at those same scales. 3. Fish and invertebrate communities were largely associated with different environmental variables, although the composition of both communities was strongly correlated with stream size across all three scales. 4. Predictive taxa‐loss models for fish assemblages were less sensitive and precise than models for invertebrate assemblages, likely because of the relatively low number of common fish taxa in our data set. Both models, however, distinguished reference from non‐reference sites. 5. The importance of concordance, geographic context and scale are discussed in relation to the design and interpretation of stream integrity indicators. In particular, our findings suggest that community concordance should not be viewed as a substitute for an evaluation of how assemblages respond to environmental stressors.     

Comparative Environmental Assessment in the Studies of Benthic Diatom,Macroinvertebrate, and Fish Communities

Community response to environmental gradients operating at hierarchical scales was assessed in studies of benthic diatoms, macroinvertebrates and fish from 44 stream sites in the New York City watershed. Hierarchical cluster analysis (TWINSPAN) of diatoms and fish partitioned the study sites into four groups, i.e., acid streams, reservoir outlets and wetland streams, large eutrophic streams, and small eutrophic streams; macroinvertebrate TWINSPAN distinguished an additional group of silty eutrophic streams. The correspondence among the three assemblage TWINSPAN groupings was moderate, ranging from 51 to 57%. The similarity across the four major group types was the highest among large eutrophic stream and acid stream assemblages, and the lowest among small eutrophic stream assemblages. Stepwise discriminant function analysis revealed that environmental factors discriminated most effectively the diatom grouping and least effectively the fish grouping. The best environmental predictors for diatom and macroinvertebrate grouping were conductance and percent surface water, while population density was most powerful in separating the fish groups. Carbaryl was the only pesticide that correlated with macroinvertebrate grouping. Partial redundancy analyses suggested a differential dependence of freshwater communities on the scale of the environmental factors to which they respond. The role of small‐scale habitat and habitatland cover/land use interaction steadily increased across the diatom, macroinvertebrate, and fish assemblages, whereas the effect of large‐scale land cover/land use declined.     

Landscape variables influence taxonomic and trait composition of insect assemblages in Neotropical savanna streams

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Termites as bioindicators of habitat quality in the caatinga, Brazil: is there agreement between structural habitat variables and the sampled assemblages?

The composition of termite assemblages was analyzed in three caatinga sites of the Esta??o Ecológica do Seridó, located in the municipality of Serra Negra do Norte, in the state of Rio Grande do Norte, Brazil. These sites have been subjected to selective logging, and cleared for pasture and farming. A standardized sampling protocol for termite assemblages (30h/person/site) was conducted between September 2007 and February 2009. At each site we measured environmental variables, such as soil pH and organic matter, necromass stock, vegetation height, stem diameter at ankle height (DAH) and the largest and the smallest crown width. Ten species of termites, belonging to eight genera and three families, were found at the three experimental sites. Four feeding groups were sampled: wood-feeders, soil-feeders, wood-soil interface feeders and leaf-feeders. The wood-feeders were dominant in number of species and number of encounters at all sites. In general, the sites were not significantly different in relation to the environmental variables measured. The same pattern was observed for termite assemblages, where no significant differences in species richness, relative abundance and taxonomic and functional composition were observed between the three sites. The agreement between composition of assemblages and environmental variables reinforces the potential of termites as biological indicators of habitat quality.     

The importance of environmental heterogeneity for species diversity and assemblage structure in Bornean stream frogs

1. Several hypotheses have been proposed to explain the structure of multi-species assemblages. Among these, abiotic environmental factors and biotic processes are often favoured. Several recent studies examining anuran communities identified environmental factors to be only of minor importance in the composition of leaf-litter and canopy assemblages in pristine forests. Instead, spatial effects and spatially structured environments were considered more important. 2. In this study, we investigated whether these findings could also be confirmed for very heterogeneous stream habitats in the primary rainforest of the Ulu Temburong National Park, Brunei Darussalam. We thus investigated anuran assemblage compositions on 50 stream sites with regard to environmental and spatial influences. 3. Cross-product correlations indicated that both factors (spatial and environmental parameters) determined assemblage composition of anurans. Environment itself may be spatially structured, yet this interrelation did not contribute to the explainable variation of frog community compositions within the study area. 4. Detailed analyses of the environmental parameters with nonmetric multidimensional scaling revealed that community structure was mostly affected by three major environmental characters: stream turbidity, river size and the density of understorey vegetation. Based on these habitat characteristics, we assigned species to three distinct habitat guilds. 5. The results underline the importance of riparian habitat heterogeneity in pristine forests in structuring anuran assemblages. We conclude that different anuran assemblages, that is, leaf litter, canopy and stream communities, follow different assemblage rules and thus are not directly comparable.     

Downstream fish assemblage response to river impoundment varies with degree of hydrologic alteration

River impoundments can fundamentally restructure downstream fish assemblages by altering flow regimes. However, the degree of alteration and associated ecological change may depend on pre-existing hydrologic regimes. We used long-term datasets to compare downstream hydrological and fish assemblage responses to impoundment in two catchments classified as having intermittent and perennial-flashy natural hydrologic regimes. We observed significant shifts in fish assemblage structure at both sites after stream impoundment. The historically intermittent stream shifted to a stable perennial flow regime. Changes in fish assemblage structure covaried with changes in five different components of the flow regime; most species that increased in abundance require fluvial habitats and likely benefited from increased flows during historically low flow seasons. Shifts in fish assemblage structure were also observed in the perennial stream, despite minimal flow alteration after impoundment; however, most species shifts were associated with lentic environments, and were more likely related to proximity of reservoirs in the drainage system rather than changes in stream flow. Findings from this study confirm that downstream fish assemblage response to river impoundment can be associated with high levels of hydrologic alteration, but other factors including expansion of lentic species into lotic environments also influence shifts in assemblage structure.     

Different roles of environmental variables and spatial factors in structuring stream benthic diatom and macroinvertebrate in Yangtze River Delta,China

Exploring the relative contribution of spatial factors and environmental variables in shaping communities is of widespread interest in biodiversity conservation and environmental management. Stream communities are hierarchically regulated by environmental variables over multiple spatial scales, and the reaction of different organisms to stressors are still equivocal. We sampled both macroinvertebrates and diatom at 80 sites and additional 10 sites for macroinvertebrates, field measured and laboratory analyzed environmental variables, from the tributaries of Qiantang River, Yangtze River Delta China in 2011. We used PCNM (principal coordinates of neighbor matrices) to generate spatial predictors. We applied redundancy analysis and variation partitioning procedures to identify key spatial and environmental factors, and to quantify their relative roles in shaping diatom and macroinvertebrate assemblages. Our results demonstrated the role of spatial and environmental variables differed in shaping benthic diatom and macroinvertebrate. Diatom assemblage variations were better explained by spatial factors, however macroinvertebrate assemblage variations were better explained by environmental variables. In terms of environmental variables, catchment scale variables (e.g., land use estimators, land use diversity) played the primary role in determining the patterns of both diatom and macroinvertebrate assemblages, whereas the influence of reach-scale variables (e.g., pH, substrates, and nutrients) appeared less. However, nutrients were the stronger factors influencing benthic diatom, whereas physical habitat (e.g., substrates) played more important role than water chemistry in structuring macroinvertebrates. Our results provided more evidence to the incorporation of spatial factors interpreting spatial patterns of stream organisms, and highlighted the useful of multiple organisms and environmental variables at different spatial scales in diagnosing mechanism of stream degradation and in building a sound stream conditions monitoring program for Yangtze River Delta.     

The influence of stream invertebrate composition at neighbouring sites on local assemblage composition

1. The composition of freshwater invertebrate assemblages at a location is determined by a range of physico‐chemical and biotic factors in the local environment, as well as larger‐scale spatial factors such as sources of recruits. We assessed the relative importance of the species composition of local neighbourhoods and proximal environmental factors on the composition of invertebrate assemblages. 2. Macroinvertebrate assemblages were sampled at 188 running‐water sites in the catchment of the River Rede, north‐east England. A total of 176 species were recorded. 3. Environmental data, in the form of 13 biotic and abiotic measurements that described stream physical structure, aquatic vegetation and water characteristics, were recorded for each site. Detrended correspondence analysis was then used to simplify nine of these stream environmental variables to create an index of stream structure. 4. The species composition of the invertebrate assemblages was related to the environmental variables, using an information theoretic approach. The impact of the species composition of neighbouring sites on each site was determined using Moran's I and autoregressive modelling techniques. 5. Species composition was primarily associated with water pH and stream structure. The importance of the species composition of neighbouring sites in determining local species assemblages differed markedly between taxa. The autoregressive component was low for Coleoptera, intermediate for Trichoptera and Plecoptera, and high for Ephemeroptera. 6. We hypothesise that the observed differences in the autoregressive component amongst these orders reflects variation in their dispersal abilities from neighbouring sites.     

Stream macroinvertebrate response to catchment urbanisation (Georgia, U.S.A.)

SUMMARY 1. The effects of catchment urbanisation on water quality were examined for 30 streams (stratified into 15, 50 and 100 km² ± 25% catchments) in the Etowah River basin, Georgia, U.S.A. We examined relationships between land cover (implying cover and use) in these catchments (e.g. urban, forest and agriculture) and macroinvertebrate assemblage attributes using several previously published indices to summarise macroinvertebrate response. Based on a priori predictions as to mechanisms of biotic impairment under changing land cover, additional measurements were made to assess geomorphology, hydrology and chemistry in each stream. 2. We found strong relationships between catchment land cover and stream biota. Taxon richness and other biotic indices that reflected good water quality were negatively related to urban land cover and positively related to forest land cover. Urban land cover alone explained 29–38% of the variation in some macroinvertebrate indices. Reduced water quality was detectable at c. >15% urban land cover. 3. Urban land cover correlated with a number of geomorphic variables such as stream bed sediment size (–) and total suspended solids (+) as well as a number of water chemistry variables including nitrogen and phosphorus concentrations (+), specific conductance (+) and turbidity (+). Biotic indices were better predicted by these reach scale variables than single, catchment scale land cover variables. Multiple regression models explained 69% of variation in total taxon richness and 78% of the variation in the Invertebrate Community Index (ICI) using phi variability, specific conductance and depth, and riffle phi, specific conductance and phi variability, respectively. 4. Indirect ordination analysis was used to describe assemblage and functional group changes among sites and corroborate which environmental variables were most important in driving differences in macroinvertebrate assemblages. The first axis in a non‐metric multidimensional scaling ordination was highly related to environmental variables (slope, specific conductance, phi variability; adj. R²=0.83) that were also important in our multiple regression models. 5. Catchment urbanisation resulted in less diverse and more tolerant stream macroinvertebrate assemblages via increased sediment transport, reduced stream bed sediment size and increased solutes. The biotic indices that were most sensitive to environmental variation were taxon richness, EPT richness and the ICI. Our results were largely consistent over the range in basin size we tested.     

钱塘江中游流域不同空间尺度环境因子对底栖动物群落的影响

溪流底栖动物群落结构受不同空间尺度环境因子的共同作用。基于2010年钱塘江中游流域60个样点的大型底栖无脊椎动物和环境变量数据,寻找与研究流域底栖动物群落结构变化密切相关的关键环境变量,解析流域尺度和河段尺度的环境因子对底栖动物群落的相对影响。PCA分析表明该区域的主要环境梯度是流域内的土地利用类型及其引起的溪流物理生境退化程度和水体营养状态。CCA分析发现影响底栖动物群落的流域尺度的关键环境变量是纬度、海拔、样点所在流域大小、森林用地百分比,河段尺度是总氮、总磷、钙浓度、二氧化硅浓度和平均底质得分。偏CCA分析得到两种尺度环境因子对底栖动物变异的总解释量为26.4%,流域尺度和河段尺度变量分别为总解释量的50%和31%;方差分解结果表明研究区域大型底栖无脊椎动物受到两种尺度环境因子的综合影响,且流域尺度环境因子较河段尺度环境因子更为重要,体现了其在溪流生态系统保护、恢复、监测和评价中的重要参考价值。     

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.     

Effects of oil palm plantations on habitat structure and fish assemblages in Amazon streams

The aim of this research is to assess the effects of oil palm plantations on stream habitat and their fish assemblage diversity. We hypothesize that streams which drain through oil palm plantations tend to be less heterogeneous, limiting the occurrence of many species, than streams that drain through forest fragments, which support higher fish diversity. A total of 17 streams were sampled; eight in forest fragments and nine in oil palm plantations. Environmental and biological variables were sampled along 150 m stretch in each stream. Of the 242 environmental variables measured, ten were considered important to assess the condition of structural habitat, and out of these variables, four were considered relevant in the distinction between streams in oil palm plantations and forest fragments. A total of 7245 fishes were collected, belonging to 63 species. Unlike our original hypothesis, the species richness did not differ between forest fragment and oil palm plantations streams, showing that it is not a good divert measure in streams disturbance assessment. However, fish assemblages differed in species composition, and 56 species were recorded in oil palm plantation streams, while 44 species were recorded in forest fragments streams. Some species were identified as indicators of either altered (Aequidens tetramerus and Apistogramma agassizii) or undisturbed areas (Helogenes marmoratus). Overall, oil palm plantations were proven to change stream habitat structure and fish species distribution, corroborating other studies that have evidenced changes in patterns of biological community structure due to impacts by different land uses.     

Local and regional determinants of stream fish assemblage structure: inferences based on taxonomic vs. functional groups

Aim  To examine the roles of local and regional environmental variables and biotic interactions in determining the structure of local stream fish assemblages, and to compare results derived from analyses based on taxonomic and functional groups.
Location  Texas, USA.
Methods  Species abundance data were compiled for 157 stream fish assemblages in several river basins across Texas. Species were condensed into functional groups based on trophic and life-history characteristics. Local and regional environmental variables were either measured at each location or determined from scale maps and public-access data bases. The original taxonomic and functional group data sets were analysed using similarity indices, null models of co-occurrence, and direct and indirect ordination techniques. Results derived from taxonomic and functional group data sets are compared.
Results  Inferences regarding the relative roles of local and larger-scale factors in determining stream fish assemblage structure differ dramatically between analyses of taxonomic and functional groups. Taxonomic analyses suggest a prominent role of regional-scale environmental factors, and local assemblages sorted according to a biogeographic pattern. Functional group analyses suggest almost equal roles of factors representative of local and larger scales, and assemblages were distinguished by a habitat template irrespective of geographic region.
Main conclusions  The structure of local stream fish assemblages is determined ultimately by factors representing multiple scales, with the relative importance of each depending on the biological unit employed (species or functional groups). We suggest that analyses using functional groups can more directly infer ecological responses to environmental variation, and therefore may provide a more fruitful avenue for developing and testing ecological theory of community organization across biogeographic scales.     

Fish ecoregions of Kansas: stream fish assemblage patterns and associated environmental correlates

Synopsis Principal components analysis was performed on fish presence/absence data for 39 common fish species from 410 stream sites in Kansas. The analysis confirmed ten ecologically meaningful fish assemblages, based on species associations. Factor scores based on these assemblages were then clustered into six geographic areas or fish ecoregions. Canonical discriminant analysis identified environmental variables that distinguished the derived fish ecoregions. Mean annual runoff, mean annual growing season, and discharge appear most important. Mean width, mean depth, chloride concentration, water temperature, substrate type, gradient, and percent of pool habitat were less important. Correspondence exists between these fish ecoregions and the patterns of physiographic regions, river basins, geology, soil, and potential natural vegetation in Kansas. The multivariate statistical approach used to classify fish ecoregions should have considerable potential value for fish assessment and management purposes in areas other than the state of Kansas.     

Spatial and temporal variation in fish assemblages of drying stream pools: The role of abiotic and biotic factors

Droughts and summer drying create unusual temporary aquatic habitats in the form of isolated pools in many small streams around the world. To examine spatial and temporal variation in fish community structure of drying stream pools, their relation to abiotic environmental variables, and associations among species, fish were sampled during summer 1995 and 1996 from pools of four streams in the Ozark mountains, Arkansas, USA. Redundancy analysis of physical-chemical variables showed significant differences among stream sites, but no significant difference between years or stream site by year interaction. Stream sites separated consistently along axes one (habitat heterogeneity) and two (temperature/canopy cover) in both years. Redundancy analysis of fish species-size class densities showed a significant stream site by year interaction. Groupings of stream sites based on fish assemblages were not well explained by physical-chemical variables measured at the pool scale, but were related to location within the drainage basin, and these groupings differed between years. There were 27 (15.8%) and 10 (5.8%) significant associations found among fish species-size classes in 1995 and 1996, respectively, and all but two significant associations in 1995 were positive. Pool depth, habitat heterogeneity, pool size and dissolved oxygen/canopy cover were important local abiotic factors depending on response variables examined. In both years, large fish total density, large central stoneroller density (80 mm TL), and small sunfish (<80 mm TL) density were positively related to pool depth. Otherwise, there was no consistent relationship between physical-chemical variables and dependent variables (fish density and species richness) within a year or between years for a given dependent variable. These results support the hypothesis that local abiotic factors are important in structuring fish assemblages in harsh environments, but the importance of those factors varies temporally, and regional influences appear to override local abiotic conditions as factors structuring fish assemblages in drying stream pools. Predation by terrestrial vertebrates may also be an important factor structuring these fish assemblages that has been largely overlooked.     

Longitudinal structure of fish assemblages in a minimally disrupted stream

The longitudinal distribution patterns of fish species are affected by both natural and anthropogenic variables. The role of these factors on the formation of species assemblages is well documented in North America and Western Europe, but detailed information is lacking from Central and Eastern Europe, and the Carpathian region especially. Therefore, we examined the structure of fish assemblages in response to six key environmental parameters in a natural stream system (Udava stream basin, Slovakia). We used the indirect ordination method of gradient analysis (Detrended Correspondence Analysis, DCA) to analyse the species groups and their connections to the sampled sites and to recognize the strongest gradient of assemblage composition. Subsequently, we used the direct ordination method (Canonical Correspondence Analysis, CCA) to identify the strongest gradients in relation to selected variables. Two major gradients were identified that follow the upstream-downstream pattern of fish communities and three variables (distance from source, depth and site slope) are correlated with the first CCA axis (P < 0.05) and two variables (depth and vegetation cover) are correlated with the second CCA axis (P < 0.05). We assume that these factors influence the temperature and the amount of dissolved oxygen that can cause oxygen and temperature stress to intolerant species (e.g., salmonids). Based on these results, we assume that the economically important species, brown trout and grayling, are not native to the stream basin and this status is only the consequence of natural factors. Furthermore, the results suggest that the Udava stream offers favourable conditions for fish species distribution — a view supported by the high variability of particular variables within the proposed model.