Intercontinental trends in functional and phylogenetic structure of stream fish assemblages

Understanding of community assembly has been improved by phylogenetic and trait‐based approaches, yet there is little consensus regarding the relative importance of alternative mechanisms and few studies have been done at large geographic and phylogenetic scales. Here, we use phylogenetic and trait dispersion approaches to determine the relative contribution of limiting similarity and environmental filtering to community assembly of stream fishes at an intercontinental scale. We sampled stream fishes from five zoogeographic regions. Analysis of traits associated with habitat use, feeding, or both resulted in more occurrences of trait underdispersion than overdispersion regardless of spatial scale or species pool. Our results suggest that environmental filtering and, to a lesser extent, species interactions were important mechanisms of community assembly for fishes inhabiting small, low‐gradient streams in all five regions. However, a large proportion of the trait dispersion values were no different from random. This suggests that stochastic factors or opposing assembly mechanisms also influenced stream fish assemblages and their trait dispersion patterns. Local assemblages tended to have lower functional diversity in microhabitats with high water velocity, shallow water depth, and homogeneous substrates lacking structural complexity, lending support for the stress‐dominance hypothesis. A high prevalence of functional underdispersion coupled with phylogenetic underdispersion could reflect phylogenetic niche conservatism and/or stabilizing selection. These findings imply that environmental filtering of stream fish assemblages is not only deterministic, but also influences assemblage structure in a fairly consistent manner worldwide.     

Convergence of temperate and tropical stream fish assemblages

The hypothesis of convergence takes the deterministic view that community (or assemblage) structure can be predicted from the environment, and that the environment is expected to drive evolution in a predictable direction. Here we present results of a comparative study of freshwater fish assemblages from headwater streams in four continents (Europe, North America, Africa and South America), with the general objective of testing whether these assemblages display convergent structures under comparable environmental conditions (i.e. assemblage position in the stream longitudinal continuum). We tested this hypothesis by comparing species richness and trophic guilds of those stream fish assemblages represented in available data from multiple sites on each continent. Independent of phylogenetic and historical constraints, fish assemblage richness and trophic structure in the four continents converged along the stream continua to a substantial degree. For the four continents, assemblage richness increased, the proportion of invertivorous species decreased, and the proportion of omnivorous species increased from upstream to downstream, supporting theoretical predictions of the river continuum concept. However, the herbivore/detritivore and piscivore guilds were virtually absent from our small European and North American stream sites, unlike our African and South American stream sites. This divergence can be linked to differences in energy availability between temperate and tropical systems.     

Gradients in stream fish assemblages across a Mediterranean landscape: contributions of environmental factors and spatial structure

1. Despite wide recognition that fish assemblages are influenced by factors operating over a range of spatial scales, little effort has been devoted to quantifying large‐scale variation and the multiscale dependencies of assemblage patterns and processes. This is particularly true for Mediterranean streams, where seasonally predictable drying‐up may lead to a strong association between assemblage attributes and large‐scale factors affecting the distribution of population sources and extinction likelihood. 2. The contribution of large‐scale factors to stream fish assemblage variation was quantified across a Mediterranean landscape, in south‐west Portugal. Fish abundance and species composition were estimated at 166 sites across third‐ to sixth‐order streams, in March–July 1998. Variance partitioning by redundancy analyses was used to analyse assemblage variation against three sets of predictor variables: environmental (catchment position, and geomorphic and hydrological factors), large‐scale spatial trends and neighbourhood effects. 3. Environmental variables and spatial trends accounted for 34.6% of the assemblage variation across the entire region, and for 36.6 and 57.8% within the two largest catchments (Mira and Seixe). Neighbourhood effects were analysed at the catchment scale, increasing the explained variation to 56.1% (Mira) and 70.7% (Seixe). 4. A prevailing environmental gradient was reflected in an increase in the abundance of all species and size‐classes in relation to catchment position, with more fish present in larger streams and in downstream reaches. Variables describing geomorphic and hydrological settings were less important in explaining assemblage variation. 5. Spatial trends always accounted for the smallest fraction of assemblage variation, and they were probably associated with historical barriers to fish dispersal. The strong neighbourhood effects may be related to spatially autocorrelated habitat conditions, but they are also a likely consequence of fish emigration/extinction and colonisation processes. 6. These results emphasise that a substantial proportion of fish assemblage variation in Mediterranean streams may be explained by large‐scale factors, irrespective of microhabitats and local biotic interactions. It is suggested that this pattern results to a large extent from the seasonal drying‐up, with the summer shortage of surface water limiting fish occurrence in headwaters, and consequently the key core areas for fish concentrating in larger streams and tributaries adjacent to large streams because of neighbourhood effects.     

Climate change hastens the turnover of stream fish assemblages

Stream fish are expected to be significantly influenced by climate change, as they are ectothermic animals whose dispersal is limited within hydrographic networks. Nonetheless, they are also controlled by other physical factors that may prevent them moving to new thermally suitable sites. Using presence–absence records in 655 sites widespread throughout nine French river units, we predicted the potential future distribution of 30 common stream fish species facing temperature warming and change in precipitation regime. We also assessed the potential impacts on fish assemblages' structure and diversity. Only cold-water species, whose diversity is very low in French streams, were predicted to experience a strong reduction in the number of suitable sites. In contrast, most cool-water and warm-water fish species were projected to colonize many newly suitable sites. Considering that cold headwater streams are the most numerous on the Earth's surface, our results suggested that headwater species would undergo a deleterious effect of climate change, whereas downstream species would expand their range by migrating to sites located in intermediate streams or upstream. As a result, local species richness was forecasted to increase greatly and high turnover rates indicated future fundamental changes in assemblages' structure. Changes in assemblage composition were also positively related to the intensity of warming. Overall, these results (1) stressed the importance of accounting for both climatic and topographic factors when assessing the future distribution of riverine fish species and (2) may be viewed as a first estimation of climate change impacts on European freshwater fish assemblages.     

Longitudinal distribution of macroinvertebrate assemblages in a glacially influenced stream system in the Italian Alps

1. The longitudinal distribution of macroinvertebrates was investigated in June, August and September 1996 and 1997 in the Conca glacial stream and its tributary (Italian Alps; 46°N, 10°E). The principal aim was to test the 22 model that predicts the succession of faunal groups downstream of the glacial snout in relation to water temperature and channel stability. The effect of a non‐glacial tributary on the taxonomic richness and density patterns occurring in the glacial stream was also considered. 2. Channel stability showed an atypical longitudinal trend in the Conca glacial stream, being high in the upper part with Pfankuch Index values between 30 and 33. Water temperature exceeded 6 °C at all stations, with average values below 2 °C occurring only within 700 m from the glacial snout. 3. Taxonomic richness and diversity increased downstream. Taxonomic richness in the glacial stream (at about 1.5 km from the glacier) was comparable with the tributary and the reach after the confluence. Abundance also increased downstream in the glacial stream, but not as greatly as the number of taxa. 4. At higher taxonomic levels, the community structure in the tributary stations appeared to be similar to the two stations in the glacial stream just upstream of the confluence. The effect of the tributary was evident mainly at the genus or species level of the Chironomidae community. Some taxa found in the non‐glacial stream (e.g. Cricotopus fuscus, Eukiefferiella coerulescens, Metriocnemus sp., Paratrichocladius rufiventris, P. skirwitensis, Rheocricotopus effusus and Smittia sp.) were found also in the Conca stream but only after the confluence. 5. The upper glacial reach (within 700 m from the glacier snout) was dominated by the chironomid Diamesa spp. Less than 400 m from the glacier other Diamesinae (Pseudokiefferiella parva) and a few Orthocladiinae, especially Orthocladius (Euorthocladius) rivicola gr., colonized the stream. Some Diamesinae maintained relatively dense populations at mean water temperature around 5 °C, while some Orthocladiinae colonized reaches with mean water temperature <3 °C. 6. Contrary to the 22 model, Dipteran families such as Empididae and Limoniidae were more abundant in the upper stations than Simuliidae; non‐insects such as Nematoda and Oligochaeta were also numerous at some sites. Leuctridae, Taeniopterygidae and Nemouridae were the first Plecoptera to appear upstream, while Chloroperlidae were restricted to the lower reaches. Among Ephemeroptera, Heptageniidae were more abundant than Baetidae in the glacial sites. 7. In this glacial system channel stability and maximum temperature did not show the expected longitudinal trend and thus a typical kryal community was confined within 700 m from the glacier snout where summer mean water temperature was below 4 °C.     

Effects of floods on fish assemblages in an intermittent prairie stream

1. Floods are major disturbances to stream ecosystems that can kill or displace organisms and modify habitats. Many studies have reported changes in fish assemblages after a single flood, but few studies have evaluated the importance of timing and intensity of floods on long‐term fish assemblage dynamics. 2. We used a 10‐year dataset to evaluate the effects of floods on fishes in Kings Creek, an intermittent prairie stream in north‐eastern, Kansas, U.S.A. Samples were collected seasonally at two perennial headwater sites (1995–2005) and one perennial downstream flowing site (1997–2005) allowing us to evaluate the effects of floods at different locations within a watershed. In addition, four surveys during 2003 and 2004 sampled 3–5 km of stream between the long‐term study sites to evaluate the use of intermittent reaches of this stream. 3. Because of higher discharge and bed scouring at the downstream site, we predicted that the fish assemblage would have lowered species richness and abundance following floods. In contrast, we expected increased species richness and abundance at headwater sites because floods increase stream connectivity and create the potential for colonisation from downstream reaches. 4. Akaike Information Criteria (AIC) was used to select among candidate regression models that predicted species richness and abundance based on Julian date, time since floods, season and physical habitat at each site. At the downstream site, AIC weightings suggested Julian date was the best predictor of fish assemblage structure, but no model explained >16% of the variation in species richness or community structure. Variation explained by Julian date was primarily attributed to a long‐term pattern of declining abundance of common species. At the headwater sites, there was not a single candidate model selected to predict total species abundance and assemblage structure. AIC weightings suggested variation in assemblage structure was associated with either Julian date or local habitat characteristics. 5. Fishes rapidly colonised isolated or dry habitats following floods. This was evidenced by the occurrence of fishes in intermittent reaches and the positive association between maximum daily discharge and colonisation events at both headwater sites. 6. Our study suggests floods allow dispersal into intermittent habitats with little or no downstream displacement of fishes. Movement of fishes among habitats during flooding highlights the importance of maintaining connectivity of stream networks of low to medium order prairie streams.     

Comparing fish assemblages and trophic ecology of permanent and intermittent reaches in a Mediterranean stream

Mediterranean streams are characterised by seasonal droughts, the frequency and intensity of which vary spatially and are expected to increase with global change. We studied the potential effects of drought and climate change on the fish assemblage and its trophic ecology in a Mediterranean stream by comparing an intermittent tributary with two more permanent neighbouring reaches. Although the three sites were dominated by the same two fish species, Mediterranean barbel (Barbus meridionalis) and chub (Squalius laietanus), the intermittent tributary had a lower overall fish density and fewer eel (Anguilla anguilla). The intermittent tributary had macroinvertebrates with lower density, smaller taxa and higher diversity. Fish in the intermittent tributary had significantly lower biomasses in their gut contents (adjusted for fish length) and more negative electivities than those in the permanent reaches, as well as significantly lower taxonomic diversity. These results indicate that there was reduced resource availability in the intermittent tributary, which resulted in significantly lower condition and gonadal weight (adjusted for length) of barbel and chub. The data obtained in this Mediterranean stream support the observation that reduced water flow may affect fish at both individual and assemblage levels.     

Geographic variation in patterns of nestedness among local stream fish assemblages in Virginia

Nestedness of faunal assemblages is a multi-scale phenomenon, potentially influenced by a variety of factors. Prior small-scale studies have found freshwater fish species assemblages to be nested along stream courses as a result of either selective colonization or extinction. However, within-stream gradients in temperature and other factors are correlated with the distributions of many fish species and may also contribute to nestedness. At a regional level, strongly nested patterns would require a consistent set of structuring mechanisms across streams, and correlation among species tolerances of the environmental factors that influence distribution. Thus, nestedness should be negatively associated with the spatial extent of the region analyzed and positively associated with elevational gradients (a correlate of temperature and other environmental factors). We examined these relationships for the freshwater fishes of Virginia. Regions were defined within a spatial hierarchy and included whole river drainages, portions of drainages within physiographic provinces, and smaller subdrainages. In most cases, nestedness was significantly stronger in regions of smaller spatial extent and in regions characterized by greater topographic relief. Analysis of hydrologic variability and patterns of faunal turnover provided no evidence that inter-annual colonization/extinction dynamics contributed to elevational differences in nestedness. These results suggest that, at regional scales, nestedness is influenced by interactions between biotic and abiotic factors, and that the strongest nestedness is likely to occur where a small number of organizational processes predominate, i.e., over small spatial extents and regions exhibiting strong environmental gradients.The Virginia Cooperate Fish and Wildlife Research Unit is jointly sponsored by United States Geological Survey, Virginia Polytechnic Institute and State University, and Virginia Department of Game and Inland Fisheries.     

Disturbance-driven changes in fish assemblages caused by a sudden increase in salinity in a perennial desert stream

Disturbance is a key factor in structuring fish assemblages in rivers; characterizing reactions to disturbance is especially important in systems containing endangered species due to their vulnerability to extinction. A natural disturbance event occurred in the Virgin River in 1985, when normal flow patterns changed due to the appearance of a sinkhole upstream of the hot saline Pah Tempe Springs. This led to abnormally elevated levels of temperature and salinity that propagated downstream. We examined changes in fish assemblages at five sites in a 21 km reach downstream from the hot springs in the mainstem Virgin River, which were monitored monthly (with the exception of 1 site monitored biannually) between 1984 and 1987. Specifically, we compared the responses of the different fish species in space and time. Of the six native species investigated, speckled dace, spinedace, and the endangered woundfin experienced declines in abundance at sites closest to the disturbance. In contrast, Virgin River chub, another endangered species, experienced increases in abundance in sites nearest the disturbance, and desert sucker and flannelmouth sucker showed no change. Redundancy analysis confirmed that changes in assemblage structure were significant. Differential responses to the disturbance appeared to correlate with differences in feeding ecology, with insectivores being the most affected group. In terms of recovery, species with higher abundances in neighboring sites experienced faster recoveries than those with lower abundances. This study characterizes disturbance reactions in the Virgin River, and emphasizes the susceptibility of rare species to natural disturbances and subsequent depression of recolonization rates.     

Spatial and temporal variation of fish assemblages in a subtropical small stream of the Huangshan Mountain

Spatial and temporal variation of fish assemblages were investigated seasonally from May 2007 to February 2008 across 11 study sites in a subtropical small stream, the Puxi Stream, of the Huangshan Mountain. Along the longitudinal gradient from headwater to downstream, fish species richness and abundance increased gradually, but then decreased significantly at the lower reaches. The highest species richness and abundance were observed in August and the lowest in February. Based on analysis of similarities (...     

The role of trait‐based approaches in understanding stream fish assemblages

1. The use of trait‐based approaches to examine the ecology of stream fish assemblages is increasing. However, selection of traits that will be useful in testing spatial or temporal hypotheses about ecological organisation is currently limited by availability of data, rather than empirical evaluation. 2. We analysed two data sets of stream fish assemblages to compare taxonomy and trait‐based approaches. The Wabash River temporal data set is based on 25 years of boat electrofishing collections over a 230‐km river distance. The Indiana Department of Environmental Management data set of stream collections in the state of Indiana was selected to represent a spatial database. We compared several trait‐based approaches: reproductive guilds, life history variables, biomonitoring metrics, ecosystem‐based functional guilds and feeding and ecosystem interaction guilds. 3. Analyses of fish assemblages that are designed to detect how environmental variation structures fish assemblages can expect similar results using taxonomic or trait‐based approaches. Results of trait‐based approaches will vary according to the spatial extent of the region and the number of unique entities of trait groups for a given data set. However, taxonomic analyses accounted for more variation than any trait‐based analyses.     

Characterizing coal and mineral mines as a regional source of stress to stream fish assemblages

Mining impacts on stream systems have historically been studied over small spatial scales, yet investigations over large areas may be useful for characterizing mining as a regional source of stress to stream fishes. The associations between co-occurring stream fish assemblages and densities of various “classes” of mining occurring in the same catchments were tested using threshold analysis. Threshold analysis identifies the point at which fish assemblages change substantially from best available habitat conditions with increasing disturbance. As this occurred over large regions, species comprising fish assemblages were represented by various functional traits as well as other measures of interest to management (characterizing reproductive ecology and life history, habitat preferences, trophic ecology, assemblage diversity and evenness, tolerance to anthropogenic disturbance and state-recognized game species). We used two threshold detection methods: change-point analysis with indicator analysis and piecewise linear regression. We accepted only those thresholds that were highly statistically significant (p ≤ 0.01) for both techniques and overlapped within ≤5% error. We found consistent, wedge-shaped declines in multiple fish metrics with increasing levels of mining in catchments, suggesting mines are a regional source of disturbance. Threshold responses were consistent across the three ecoregions occurring at low mine densities. For 47.2% of the significant thresholds, a density of only ≤0.01 mines/km² caused a threshold response. In fact, at least 25% of streams in each of our three study ecoregions have mine densities in their catchments with the potential to affect fish assemblages. Compared to other anthropogenic impacts assessed over large areas (agriculture, impervious surface or urban land use), mining had a more pronounced and consistent impact on fish assemblages.     

Temporal changes in replicated experimental stream fish assemblages: predictable or not?

Summary 1. Natural aquatic communities or habitats cannot be fully replicated in the wild, so little is known about how initially identical communities might change over time, or the extent to which observed changes in community structure are caused by internal factors (such as interspecific interactions or traits of individual species) versus factors external to the local community (such as abiotic disturbances or invasions of new species).
2. We quantified changes in seven initially identical fish assemblages, in habitats that were as similar as possible, in seminatural artificial streams in a 388-day trial (May 1998 to May 1999), and compared the change to that in fish assemblages in small pools of a natural stream during a year. The experimental design excluded floods, droughts, immigration or emigration. The experimental fish communities diverged significantly in composition and exhibited dissimilar trajectories in multivariate species space. Divergence among the assemblages increased from May through August, but not thereafter.
3. Differences among the experimental assemblages were influenced by differences that developed during the year in algae cover and in potential predation (due to differential survival of sunfish among units).
4. In the natural stream, fish assemblages in small pools changed more than those in the experimental units, suggesting that in natural assemblages external factors exacerbated temporal variation.
5. Our finding that initially identical assemblages, isolated from most external factors, would diverge in the structure of fish assemblages over time suggests a lack of strong internal, deterministic controls in the assemblages, and that idiosyncratic or stochastic components (chance encounters among species; vagaries in changes in the local habitat) even within habitat patches can play an important role in assemblage structure in natural systems.     

Temporal variation in foraging group structure of a size-structured stream fish community

Temporal variation in foraging group structure of a fish assemblage was examined in a flood-prone stream in southern Hokkaido, Japan. Foraging behaviour was observed underwater for four species which inhabit the water column: ayu, Plecoglossus altivelis, white-spotted charr, Salvelinus leucomaenis, masu salmon, Oncorhynchus masou, and Japanese dace, Tribolodon hakonensis, with each species being categorized into five size classes (species-size group; SSG). Based on foraging behaviour, each SSG of the fish assemblage was classified into one of four foraging groups: algae grazers, drift foragers, benthos-drift foragers, and omnivores, defined as SSG exhibiting similar foraging behaviour. All size classes of ayu, and of charr and salmon were categorized as algae grazers and drift foragers, respectively, throughout the study period. In contrast, size classes of dace were categorized as drift foragers, benthos-drift foragers, or omnivores with the same size classes often assigned to different foraging groups from month to month. Digestive tract contents of the fishes in the four foraging groups reflected their observed foraging behaviour, and foraging groups were therefore regarded as representing trophic groups. Abundance and membership of each foraging group varied in accordance with changes in abundance of SSG due to their growth, immigration, emigration, and/or mortality. Moreover, due to numerical dominance within the assemblage, plasticity in foraging behaviour of small- and medium-sized dace also played a key role in determining variability in the foraging group structure. Relative frequencies of two types of foraging behaviour, algae nipping and benthos foraging, of the small-sized dace were significantly correlated with the level of each resource, whereas no significant relationship was detected between foraging frequencies of the medium-sized dace and either resource. Fluctuations in foraging group structure within this assemblage occurred through niche shifts of some component members and by changes in SSG composition.     

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.     

Trait-based metrics as bioindicators: Responses of stream fish assemblages to a gradient of environmental degradation

The development of biomonitoring tools is increasingly appealing in light of the increasing degradation of aquatic ecosystems. In this context, we investigated the responses of stream fish communities to the gradient of environmental degradation in different basins using a variety of indices based on functional diversity, functional composition, and taxonomic diversity. We used datasets from three Brazilian ecoregions. In order to describe the gradient of environmental degradation we scored streams based on local and landscape variables. The functional structure of the assemblages was described in relation to seven functional traits related to habitat use, swimming capacity, and maneuverability. We described assemblages in each ecoregion separately in relation to 20 potential indicators, grouped in four families: indices of functional diversity based on presence/absence (FD_p/a); indices of functional diversity weighted by abundance (FD_abund); indices of functional composition (mT); and, indices based on taxonomic diversity (TDiv). The relationship between the indicators and the gradient of environmental degradation were evaluated using linear regression. We found a significant interaction effect (ANOVA, p = 0.006) between group of indices and ecoregions for the performance of indicators, suggesting a context-dependent response. The indices, on an individual basis, had variable performance and consistency among ecoregions. Four mT indicators demonstrated the highest average performance and consistency. Taxonomic diversity indicators consistently had the lowest average performance, while FD_p/a and FD_abund indicators had low average performance and variable inter-regional consistency. The differential inter-regional performance of indicators was due to the differences in the lengths of the gradients of environmental degradation. Our results indicate that functional traits have greater predictive power compared to taxonomic indicators for fish responses to a gradient of environmental degradation. Although indicators of functional composition are the most promising, we emphasize that caution is needed when generalizing functional diversity indicators across ecoregions because most of them are context-dependent.