What governs the functional diversity patterns of fishes in the headwater streams of the humid forest enclaves: environmental conditions,taxonomic diversity or biotic interactions?

The relationship between functional and taxonomic diversity is a major issue in ecology. Biodiversity in aquatic environments is strongly influenced by environmental gradients that act as dispersion and niche barriers. Environmental conditions act as filters to select functional traits, but biotic interactions also play a role in assemblage structure. In headwater streams, the relationship between functional and taxonomic diversity remains unclear. In this study we investigated how environmental conditions, taxonomic diversity and biotic interactions influence the spatial distribution of traits and functional diversity in stream fish species. Standardized sampling of fish species was carried out in 50 m sections of 16 streams located in rainforest enclaves in a semiarid region of Brazil (Caatinga biome). The functional diversity indices displayed different responses to the predictor variables used. Functional richness was mainly influenced by environmental conditions, while functional evenness was mostly determined by taxonomic diversity. On the other hand, functional dispersion was explained by a combination of environmental conditions and taxonomic diversity. The spatial distribution of fish species with the same functional traits was random, indicating that biotic interactions are not a strong predictor in these ecosystems. Channel width, pH and substrate were the most important variables in the spatial distribution of the functional traits of the fish species. Our results suggest that the functional structure of fish assemblages in headwater streams depends mainly on environmental conditions and taxonomic diversity.     

Fish assemblage convergence along stream environmental gradients: an intercontinental analysis

Species that pass through similar environmental filters, regardless of geographic proximity or evolutionary history, are expected to share many traits, resulting in similar assemblage trait distributions. Convergence of assemblage trait distributions among different biotic regions would indicate that consistent ecological processes produce repeated patterns of adaptive evolution. This study analyzes trait–environment relationships across multiple stream fish assemblages representing evolutionarily divergent faunas. We hypothesized that trait–environment patterns converge across regional faunas in response to a common set of environmental filters acting on functional traits. One hundred and ninety‐seven species and forty streams were sampled from five regions: Belize, Benin, Brazil, Cambodia and USA. By examining trait–environment plots, multiple congruent trait–environment patterns were found across all regions, indicative of a consistent set of environmental filters acting on local community assembly. The consistency of these patterns strongly suggests that water velocity and habitat structural complexity function as universal environmental filters, producing similar assemblage trait distributions in streams across all regions. Bivariate relationships were not universal, and only one of the associations between a single functional trait and single environmental variable was statistically significant across all five regions. Strong phylogenetic signal was found in traits and habitat use, which implies that niche conservatism also influenced assemblage trait distributions. Overall, results support the idea that habitat templates structure trait distributions of stream fish assemblages and do so in a consistent manner.     

Functional diversity and trait–environment relationships of stream fish assemblages in a large tropical catchment

1. The species composition of stream fish assemblages changes across the longitudinal fluvial gradient of large river basins. These changes may reflect both zonation in species distributions and environmental filtering of fish traits as stream environments change from the uplands to the lowlands of large catchments. Previous research has shown that taxonomic diversity generally increases in larger, lowland streams, and the River Continuum Concept, the River Habitat Template and other frameworks have provided expectations for what functional groups of fishes should predominate in certain stream types. However, studies addressing the functional trait composition of fish assemblages across large regions are lacking, particularly in tropical river basins. 2. We examined functional trait–environment relationships and functional diversity of stream fish assemblages in the Río Grijalva Basin in southern Mexico. Traits linked to feeding, locomotion and life history strategy were measured in fishes from streams throughout the catchment, from highland headwaters to broad, lowland streams. Relationships between functional traits and environmental variables at local and landscape scales were examined using multivariate ordination, and the convex hull volume of trait space occupied by fish assemblages was calculated as a measure of functional diversity. 3. Although there were a few exceptions, functional diversity of assemblages increased with species richness along the gradient from uplands to lowlands within the Grijalva Basin. Traits related to swimming, habitat preference and food resource use were associated with both local (e.g. substratum type, pool availability) and landscape‐scale (e.g. forest cover) environmental variables. 4. Along with taxonomic structure and diversity, the functional composition of fish assemblages changed across the longitudinal fluvial gradient of the basin. Trait–environment relationships documented in this study partially confirmed theoretical expectations and revealed patterns that may help in developing a better understanding of general functional responses of fish assemblages to environmental change.     

Geomorphology variables predict fish assemblages for forested and endorheic rivers of two continents

Stream fishes are restricted to specific environments with appropriate habitats for feeding and reproduction. Interactions between streams and surrounding landscapes influence the availability and type of fish habitat, nutrient concentrations, suspended solids, and substrate composition. Valley width and gradient are geomorphological variables that influence the frequency and intensity that a stream interacts with the surrounding landscape. For example, in constrained valleys, canyon walls are steeply sloped and valleys are narrow, limiting the movement of water into riparian zones. Wide valleys have long, flat floodplains that are inundated with high discharge. We tested for differences in fish assemblages with geomorphology variation among stream sites. We selected rivers in similar forested and endorheic ecoregion types of the United States and Mongolia. Sites where we collected were defined as geomorphologically unique river segments (i.e., functional process zones; FPZs) using an automated ArcGIS‐based tool. This tool extracts geomorphic variables at the valley and catchment scales and uses them to cluster stream segments based on their similarity. We collected a representative fish sample from replicates of FPZs. Then, we used constrained ordinations to determine whether river geomorphology could predict fish assemblage variation. Our constrained ordination approach using geomorphology to predict fish assemblages resulted in significance using fish taxonomy and traits in several watersheds. The watersheds where constrained ordinations were not successful were next analyzed with unconstrained ordinations to examine patterns among fish taxonomy and traits with geomorphology variables. Common geomorphology variables as predictors for taxonomic fish assemblages were river gradient, valley width, and valley slope. Significant geomorphology predictors of functional traits were valley width‐to‐floor width ratio, elevation, gradient, and channel sinuosity. These results provide evidence that fish assemblages respond similarly and strongly to geomorphic variables on two continents.     

Global warming and potential shift in reference conditions: the case of functional fish-based metrics

The reference condition approach, advocated by the Water Framework Directive, is the basis of most currently used multimetric indices using functional traits of fish species. The ecological status of streams is assessed by measuring the deviation of the observed trait values from the theoretical values of reference conditions in the absence of anthropogenic disturbances. While reference conditions serve as baselines for ecological assessment, they vary with natural environmental conditions. Therefore, global warming appears to be a major threat to the use of current indices for diagnosing future stream conditions, as climate change is projected to modify assemblage composition, suggesting that the functional structure of fish assemblages will also be affected. The main objectives of this study are to assess the potential effect of climate change on the trait composition of fish assemblages and the consequences for the establishment of reference conditions. The results highlight the relation between environmental, especially climatic, conditions and functional traits and project the effects of climate change on trait composition. Traits based on species intolerance are expected to be most negatively affected by the projected climatic shift. The consequences for the development of multimetric indices based on fish functional traits are discussed.     

Organization patterns of headwater-stream fish communities in the Upper Paraguay–Paraná basins

We compared environmental influences on the assemblages of stream fishes of contiguous but biogeographically distinct parts of the Upper Paraguay and Paraná basins in Brazil, aiming to describe the main distribution patterns of fish species in the headwater streams. We analyzed bimonthly samples in 10 streams in each basin from January to November 2004. Sixty fish species were collected, including 40 species in streams of the Paraguay Basin and 42 species in streams of the Paraná Basin. The species abundance rank did not differ between the basins. We found a clear differentiation between the Paraguay and Paraná stream fish assemblages. There were significant differences in species composition among streams, but no seasonal differences in species composition. Connectivity between streams explains the differences in species composition for the Paraguay streams, but not for the Paraná streams. Hydrological characteristics were the main factors determining species distribution in both basins, suggesting that the migratory capacity of each species and biogeographical barriers may act to filter the local species composition from the total species pool of colonizers. Handling editor: S. M. Thomaz     

A continental‐scale analysis of fish assemblage functional structure in European rivers

The theory of traits (life‐history, ecological and biological traits) states that a species’ characteristics might enable its persistence and development in given environmental conditions. If environment is the major factor controlling functional assemblage structure, species with similar attributes are expected to inhabit a similar environment. This study uses trait states in 849 European riverine fish assemblages to analyze the influence of environment, phylogeny and biogeography on the functional structure of these assemblages. European fish assemblages were highly structured and two main syndromes (a suite of coevolved traits) were observed: 1) assemblages dominated by stenothermal intolerant individuals and 2) assemblages dominated by eurythermal, eurytopic and tolerant individuals. Temperature and stream physical structure were the two main environmental factors explaining the diversity of fish assemblage functional structures, while the influence of biogeographic factors was weak, once environment was taken into account. This suggests that, whatever the regional species pool, similar assemblage functional structures will be found in similar environmental conditions. The phylogenetic relatedness between species might also explain to some extent the associations between the species traits observed among European fish assemblages.     

Fish assemblages in temporary ponds adjacent to 'terra-firme' streams in Central Amazonia

1. The effect of habitat structural features and physicochemical characteristics of the water on the composition and richness of fish assemblages in temporary ponds near streams were examined at three spatial scales: among ponds, among streams and between drainage basins, in a ‘terra‐firme’ (not subject to long‐term flooding) forest reserve in Central Amazonia. 2. The fish assemblage in temporary ponds was composed of subsets of 18 small‐bodied species widely distributed in the reserve. The assemblages had a nested subset structure, where smaller ponds contained subgroups of the species found in larger ponds. 3. Species composition and richness in temporary ponds were similar between drainage basins, although the fish assemblages in streams differed between basins. 4. Fish assemblage structure was influenced by local factors related to habitat structure, such as pond area and depth, canopy cover and hydroperiod. Physicochemical characteristics of the water in the ponds were similar between drainage basins and had little detectable effect on the structure of pond fish assemblages. 5. No correspondence was found between the composition, richness or abundance of fishes in the ponds and in stretches of the streams adjacent to the ponds. Therefore, it is not possible to predict the composition of these temporary pond fish assemblages from the fish assemblages found in adjacent streams.     

Relationships between fishes and habitat in rainforest streams in Sabah, Malaysia

Distinct fish assemblages were found at the mesohabitat scale in 14 streams in eastern Sabah, Malaysia. Sites were designated a priori as pool, run or riffle on the basis of physical habitat structure and properties. Principal components analysis of physical habitat data confirmed the validity of the a priori designation with a major axis of three correlated variables: water velocity, depth and substratum type. Canonical discriminant analysis on fish abundance and biomass data confirmed the existence of a specialized assemblage of fishes from riffle areas of all streams. Overall, pool and run assemblages were highly variable, dependent on stream size, but also variable between streams of the same size. Multiple regression of species richness, diversity, abundance and biomass data on principal components revealed significant but low correlations with measured habitat variables. Riffle habitats showed lower species richness and diversity but high abundance. The fish assemblage in riffles was dominated by balitorid species, specialized for fast-water conditions. Pool assemblages had the highest species diversity and were dominated by cyprinid species of a number of morphological and ecological guilds. Run assemblages were intermediate in assemblage characteristics between riffle and pool assemblages. Between-stream variation in assemblage composition was less than within-stream variation. Of 38 species collected, seven could be designated as riffle specialists, 18 as pool specialists and 13 as ubiquitous, although most of the latter showed size-specific habitat use with larger size classes found in slower, deeper water.     

Ecological values of Hamilton urban streams (North Island, New Zealand): constraints and opportunities for restoration

Urban streams globally are characterised by degraded habitat conditions and low aquatic biodiversity, but are increasingly becoming the focus of restoration activities. We investigated habitat quality, ecological function, and fish and macroinvertebrate community composition of gully streams in Hamilton City, New Zealand, and compared these with a selection of periurban sites surrounded by rural land. A similar complement of fish species was found at urban and periurban sites, including two threatened species, with only one introduced fish widespread (Gambusia affinis). Stream macroinvertebrate community metrics indicated low ecological condition at most urban and periurban sites, but highlighted the presence of one high value urban site with a fauna dominated by sensitive taxa. Light-trapping around seepages in city gullies revealed the presence of several caddisfly species normally associated with native forest, suggesting that seepage habitats can provide important refugia for some aquatic insects in urban environments. Qualitative measures of stream habitat were not significantly different between urban and periurban sites, but urban streams had significantly lower hydraulic function and higher biogeochemical function than periurban streams. These functional differences are thought to reflect, respectively, (1) the combined effects of channel modification and stormwater hydrology, and (2) the influence of riparian vegetation providing shade and enhancing habitat in streams. Significant relationships between some macroinvertebrate community metrics and riparian vegetation buffering and bank protection suggest that riparian enhancement may have beneficial ecological outcomes in some urban streams. Other actions that may contribute to urban stream restoration goals include an integrated catchment approach to resolving fish passage issues, active reintroduction of wood to streams to enhance cover and habitat heterogeneity, and seeding of depauperate streams with native migratory fish to help initiate natural recolonisation.     

Seasonal variation in the composition of fish assemblages in small Amazonian forest streams: evidence for predictable changes

1. The effects of seasonal inundation on the biology of fishes on floodplains of large Amazonian rivers are well studied. However, the small seasonal changes in headwater streams are generally considered to have little effect on fish assemblages. 2. In this study, we analysed seasonal changes in the species composition and abundance of fish in small Amazonian forest streams. We sampled fish with hand and seine nets in headwater streams in a 10 000 ha terra‐firme forest reserve near Manaus, Brazil. Each stream was surveyed at the end of the 2005 dry season, at the beginning of the 2006 rainy season and at the beginning of the 2006 dry season, by means of a standardized sampling effort. 3. The numbers of individuals and species caught were higher in the dry season, but rarefaction analyses indicated that greater species numbers could have been due simply to the larger number of individuals caught. 4. Between the dry and rainy season, the direction of changes in species composition in multivariate space varied among sites, especially for quantitative (abundance) data. However, the observed variation among sites was the less than expected if the directions of change were random. 5. Fish assemblages in the second dry season were more similar to those in the previous dry season than expected if changes in species composition among seasons were random. This indicates that a general seasonal pattern in fish assemblages can be detected, despite the existence of some erratic site‐specific changes. 6. Most of the species that showed large seasonal variations in density occupy temporary ponds during the rainy season, when much of the valley is inundated and pond networks form adjacent to streams. Short‐duration lateral migrations to these ponds may play an important role in the seasonal fish‐assemblage dynamics in Amazonian headwater streams. 7. Our results contrast with previous studies on small Amazonian streams, which have found little seasonal change in fish assemblages, and highlight the importance of the abundance of common species as an indicator of general fish assemblage structure in biological monitoring programmes.     

Seasonal fluctuations in taxonomic and functional diversity in assemblages of catfishes in the Venezuelan Arauca River Floodplain

The Arauca River harbors a considerable fish biodiversity, yet it remains poorly sampled and the knowledge about its fish communities is incomplete. We studied the taxonomic and functional composition and diversity of catfish assemblages in one lotic and one lentic habitat of a segment of this floodplain during one hydrological cycle, from August 2014 to August 2015, comprising six samplings using gillnets, and identified the relationships between catfish traits and the environment based on five limnological variables. The functional structure was studied through 11 morphological measures and the species diets. We identified 31 catfish species and nine trophic groups. There were significant differences in species composition and diversity between habitats and months, but the functional structure of the catfishes was only determined by the temporal variability. The functional structure of the catfish assemblages was ordered by morphological traits related to locomotion, habitat use, and trophic ecology and these traits were correlated with the limnological variables, supporting the existence of environmental filters. Integrating the taxonomic and functional perspectives can provide a more complete picture of the ecological patterns of freshwater fish assemblages in the Neotropics and can help us predict how these assemblages might change in response to anthropogenic alterations of the environment.     

河流鱼类分类群和功能群的纵向梯度格局——以新安江流域为例

确定鱼类群落的空间格局是保护和管理河流鱼类多样性的基础。尽管河流鱼类分类群（基于物种组成）的纵向梯度格局已得到大量报道,但其功能群（基于功能特征）的空间格局研究较少。以皖南山区新安江为研究流域,沿其"正源-下游"梯度共设置27个调查样点,分别于2017年5月和10月完成2次调查取样,着重研究了鱼类分类群和功能群结构的纵向梯度格局及其形成机制。共采集鱼类44种,可分为5个运动功能群和4个营养功能群,构成14个"营养-运动"复合功能群。双因素交互相似性分析结果显示,鱼类分类群和功能群均随河流级别显著变化,但两者均无显著的季节变化;根据相似性百分比分析,由1级至3级河流,数量优势物种和功能群的空间变化主要呈嵌套格局,而由3级至5级河流其变化主要呈周转格局。方差分解结果显示,局域栖息地、陆地景观和支流空间位置3类解释变量对分类群和功能群空间变化的解释率分别为33.6%和38.5%,其中,分类群受局域栖息地和支流空间位置变量的显著影响,而功能群受局域栖息地和陆地景观变量的显著影响。研究表明,沿着新安江的"上游-下游"纵向梯度,鱼类分类群和功能群的空间格局基本一致,但两者的形成机制不同：分类群的纵向梯度变化受环境过滤和扩散过程的联合影响,而功能群则主要受环境过滤影响。     

Hierarchical spatial structure of stream fish colonization and extinction

Spatial variation in extinction and colonization is expected to influence community composition over time. In stream fish communities, local species richness (alpha diversity) and species turnover (beta diversity) are thought to be regulated by high extinction rates in headwater streams and high colonization rates in downstream areas. We evaluated the spatiotemporal structure of fish communities in streams originally surveyed by Burton and Odum 1945 (Ecology 26: 182–194) in Virginia, USA and explored the effects of species traits on extinction and colonization dynamics. We documented dramatic changes in fish community structure at both the site and stream scales. Of the 34 fish species observed, 20 (59%) were present in both time periods, but 11 (32%) colonized the study area and three (9%) were extirpated over time. Within streams, alpha diversity increased in two of three streams but beta diversity decreased dramatically in all streams due to fish community homogenization caused by colonization of common species and extirpation of rare species. Among streams, however, fish communities differentiated over time. Regression trees indicated that reproductive life‐history traits such as spawning mound construction, associations with mound‐building species, and high fecundity were important predictors of species persistence or colonization. Conversely, native fishes not associated with mound‐building exhibited the highest rates of extirpation from streams. Our results demonstrate that stream fish colonization and extinction dynamics exhibit hierarchical spatial structure and suggest that mound‐building fishes serve as keystone species for colonization of headwater streams.     

Biogeography,ecoregions, and geomorphology affect fish species composition in streams of eastern Oklahoma,USA

Stream fish assemblages are structured by biogeographical, physical and biological factors acting on different spatial scales. We determined how physical factors, geomorphology and stream habitat, influenced fish species composition (presence–absence) in eastern Oklahoma, USA relative to the ecoregion and biogeographic effects previously reported. We sampled fish assemblages and surveyed geomorphology and habitat at 107 stream sites in the Boston Mountains, Ouachita Mountains, and Ozark Highlands ecoregions in eastern Oklahoma. Partial canonical correspondence analyses (pCCAs) and variance partitioning showed that patterns of endemism related to drainage basins and ecoregions explained important variation in fish species composition in all streams, but stream size and local channel morphology explained more variation overall. Stream size effects were most important in explaining variability in fish species composition in both northeastern and southeastern Oklahoma streams. Local channel morphology and substrate characteristics were secondarily important. Variables typically considered important as fish habitat (aquatic vegetation, etc.) had little effect on fish species composition.     

Watershed urbanization alters the composition and function of stream bacterial communities

Watershed urbanization leads to dramatic changes in draining streams, with urban streams receiving a high frequency of scouring flows, together with the nutrient, contaminant, and thermal pollution associated with urbanization. These changes are known to cause significant losses of sensitive insect and fish species from urban streams, yet little is known about how these changes affect the composition and function of stream microbial communities. Over the course of two years, we repeatedly sampled sediments from eight central North Carolina streams affected to varying degrees by watershed urbanization. For each stream and sampling date, we characterized both overall and denitrifying bacterial communities and measured denitrification potentials. Denitrification is an ecologically important process, mediated by denitrifying bacteria that use nitrate and organic carbon as substrates. Differences in overall and denitrifying bacterial community composition were strongly associated with the gradient in urbanization. Denitrification potentials, which varied widely, were not significantly associated with substrate supply. By incorporating information on the community composition of denitrifying bacteria together with substrate supply in a linear mixed-effects model, we explained 45% of the variation in denitrification potential (p-value<0.001). Our results suggest that (1) the composition of stream bacterial communities change in response to watershed urbanization and (2) such changes may have important consequences for critical ecosystem functions such as denitrification.     

Effects of pasture conversion to sugarcane for biofuel production on stream fish assemblages in tropical agroecosystems

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Fishes of Indiana streams: current and historic assemblage structure

We examined Indiana fish assemblages using taxonomy and ecological categories to assess temporal shifts in community structure and recent environmental relationships. Historic (1945) and recent (1996–2007) presence/absence data were compiled by subbasin and analyzed with Nonmetric Multidimensional Scaling (NMS) ordination and by species richness. Canonical Correspondence Analysis (CCA) was used to test taxonomic identity and ecological category abundance data for explanation with recent (1996–2007) environmental variables. We found a decrease in assemblage heterogeneity for recent assemblages and an increase in the number of tolerant species per subbasin. Recent Indiana streams are dominated by tolerant fishes with generalist life history strategies and low functional variation. The use of ecological categories resulted in weaker relationships with environmental variables than analyses with taxonomic identities. Analyses using taxonomy resulted in strong assemblage explanation from stream size and flow variation, while analyses using ecological categories resulted in strong assemblage explanation from habitat variation in silt substrates and flow. Analyses of recent assemblage structure using ecological categories resulted in decreased assemblage variation among subbasins than in analyses using taxonomic identities. We found that fish assemblages of Indiana streams are structured primarily by habitat complexity and have been altered during the past 50 years through multiple disturbances including fragmentation, siltation, and species introductions.     

Effects of urbanization on stream benthic invertebrate communities in Central Amazon

Urbanization and its physical and chemical effects on aquatic environments influence invertebrate communities negatively. Yet, it is not clear how urbanization affects inter-annual variation of invertebrate assemblages in streams. We 1) evaluated urbanization effects on the ecological conditions (biotic and abiotic) of streams in Manaus and 2) analyzed invertebrate community variation over time (between 2003 and 2010). Data on abiotic variables and invertebrates from 2003 were obtained from a previous study. In 2010 we sampled abiotic variables and invertebrate communities in the same low-order urban streams sampled in 2003 (n = 40). We recorded high values of total nitrogen, total phosphorous, deforestation, total impervious area (TIA), water temperature, pH, and electrical conductivity in the most urbanized streams, as compared to the least-impacted ones. In contrast, the least-impacted streams had high dissolved oxygen concentrations. Water quality was poorer in 2010 than in 2003: oxygen concentration was lower and total nitrogen, total phosphorous, deforestation, and TIA significantly higher in 2010. We recorded higher inter-annual variation of abiotic variables in the most-impacted streams as compared to the least-impacted streams. EPT (%, Ephemeroptera, Plecoptera, and Trichoptera) and richness metrics decreased with urbanization. On the other hand, % OP (percent of Oligochaeta and Psychodidae) increased with urbanization. Observed and EPT richness and% OP increased between 2003 and 2010. On the other hand, rarefied richness decreased between years. Increases of observed and EPT richness between 2003 and 2010 were related to low inter-annual variability in streams conditions; however, differences of% OP and rarefied richness were not related to inter-annual variability in environmental conditions. The degree of urbanization did not explain the magnitude of the within-stream difference of invertebrate communities between 2003 and 2010. The increased effects of urbanization represented by the abiotic variables sampled and the reduction of invertebrate richness and increased dominance of tolerant taxa indicate that public policy is not enough to protect or mitigate human impacts on the urban water systems under study.