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.     

Structure and dynamics of stream fish communities in the flood zone of the lower Purus River,Amazonas State,Brazil

The structure and dynamics of fish communities were characterized by richness, abundance, diversity and stability, during high-water and low-water periods. These analyses were performed on data from the flood zone of four streams in the lower Purus River, in the Brazilian Amazon. A total of 188 species of fish were collected, distributed among 29 families and eight orders. The statistical test showed a difference in community diversity between periods. The high-water period showed higher evenness in comparison to the low-water season. The low-water period was marked by high species abundance. A great variation in community composition between the flood and low-water periods was encountered. The adjustments for species abundance models suggested that stochastic events structure the communities. Most of the species showed a temporal variation of abundance indicating low community stability. Changes in the physico-chemical conditions of the water caused by the seasonal hydrological regime may be influencing the structuring of the fish communities.     

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.     

IS TEMPORAL OCCURRENCE OF DIATOMS RELATED TO SPECIES TRAITS,LOCAL ABUNDANCE,AND REGIONAL DISTRIBUTION?1

Water quality in streams typically changes fast, and sensitive biological indicators are crucial for monitoring these changes in water quality. Diatoms are widely used in biological stream quality assessment. However, interannual and intra‐annual variation of diatom population densities is large, often hampering a reliable quality assessment. We studied the importance of different species traits on temporal occurrence of diatom species. We also examined whether temporal occurrence of diatoms is related to species' local abundance or regional distribution using a data set collected in Finnish streams. According to the general linear model (GLM), temporal occurrence of diatoms increased with increasing local abundance and regional distribution. Species that occurred more frequently also had larger niche breadths and nonmarginal niche positions. In addition, cell size and attachment ability were positively related to species temporal occurrence. Our results imply that abundant and widely distributed species with large niches and ability to attach sustain persistent populations in varying environmental conditions typical for streams. We suggest that future studies could concentrate on monitoring these common (abundant) species when detecting the possible changes in the biological state of streams. We advise, however, to consider a relatively large number of species as many of the most common species may have low indicator values.     

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.     

Convergence of fish communities from the littoral zone of reservoirs

1. Understanding factors that regulate the assembly of communities is a main focus of ecology. Human‐engineered habitats, such as reservoirs, may provide insight into these assembly processes because they represent novel habitats that are subjected to colonization by fishes from the surrounding river basin or transported by humans. By contrasting community similarity within and among reservoirs from different drainage basins to nearby stream communities, we can test the relative constraints of reservoir habitats and regional species pools in determining species composition of reservoirs. 2. We used a large spatial database that included intensive collections from 143 stream and 28 reservoir sites within three major river basins in the Great Plains, U.S.A., to compare patterns of species diversity and community structure between streams and reservoirs and to characterize variation in fish community structure within and among major drainage basins. We expected reservoir fish faunas to reflect the regional species pool, but would be more homogeneous that stream communities because similar species are stocked and thrive in reservoirs (e.g. planktivores and piscivores), and they lack obligate stream organisms that are not shared among regional species pools. 3. We found that fish communities from reservoirs were a subset of fishes collected from streams and dominant taxa had ecological traits that would be favoured in lentic environments. Although there were regional differences in reservoir fish communities, species richness, patterns of rank abundance and community structure in reservoir communities were more homogonous across three major drainage basins than for stream communities. 4. The general pattern of convergence of reservoir fish community structure suggests their assembly is constrained by local factors such as habitat and biotic interactions, and facilitated by the introduction of species among basins. Because there is a reciprocal transfer of biota between reservoirs and streams, understanding factors structuring both habitats is necessary to evaluate the long‐term dynamics of impounded river networks.     

BIODIVERSITY RESEARCH: Native and introduced fish species richness in Mediterranean streams: the role of multiple landscape influences

Aim To examine the role of multiple landscape factors on the species richness patterns of native and introduced freshwater fish. Location Mediterranean streams, south‐western Iberian Peninsula, Europe (c. 87,000 km²). Methods We used a dataset of fish occurrences from 436 stream sites. We quantified the incremental explanatory power of multiple landscape factors in native, introduced, and overall local species richness using regression analysis. First, we related variation in local species richness across river basins to regional species richness (here, the basin species pool), area and factors of climate and topography. Second, we related within‐river basin local species richness to site’s climate and topography, and spatial structure derived from Principal Coordinates of Neighbour Matrices approach, after testing for species richness spatial autocorrelation; predicted local richness was mapped. Results Patterns of local species richness across river basins were strongly associated with regional species richness for overall, native and introduced species; annual rainfall showed a significant incremental contribution to variation in introduced species richness only. Within river basins, environmental factors were associated with local richness for the three species groups, though their contributions to the total explained variation were inferior to those of spatial factors; rainfall seasonality and stream slope were the most consistent environmental correlates for all species groups, while the influence of spatial factors was most prevalent for native species. Main conclusions Landscape factors operating among and within river basins seem to play a relevant role in shaping local species richness of both native and introduced species, and may be contingent on basin‐specific contexts. Nevertheless, local factors, such as habitat characteristics and biotic interactions and human‐induced disturbances may also be at play. Multiscale approaches incorporating a multitude of factors are strongly encouraged to facilitate a deeper understanding of the biodiversity patterns of Mediterranean streams, and to promote more effective conservation and management strategies.     

Associations between fish assemblages and environmental factors for Mediterranean-type rivers during summer

The structure of summer fish assemblages was examined along longitudinal gradients in 31 Mediterranean-type rivers of the middle Guadiana basin (south-west Iberian Peninsula), using data from 157 sites including small streams to deep rivers. An ordination analysis, based on 16 variables, was applied to species presence, using principal component and canonical correspondence analysis. The results for the habitat data were compared with those for the biological data using a Mantel analysis, and the agreement was highly significant. Spatial structure was considered by partitioning the total variability among the environmental and geographical variables. The fish assemblages showed longitudinal zoning during the summer, with species distributed over gradients of habitat size (depth), water quality (current and physico-chemical variables), and cover (substratum and vegetation), according to their adult size and life history. The size of the habitat that remained available in summer had the greatest biological effect, being the most important factor explaining fish species distribution and assemblage structure during this stressing season. Strictly spatial variation was low, but there was still a high residual variation. Habitat associations and life-history strategies are discussed for native and exotic species.     

A multi-scale comparison of trait linkages to environmental and spatial variables in fish communities across a large freshwater lake

Species present in communities are affected by the prevailing environmental conditions, and the traits that these species display may be sensitive indicators of community responses to environmental change. However, interpretation of community responses may be confounded by environmental variation at different spatial scales. Using a hierarchical approach, we assessed the spatial and temporal variation of traits in coastal fish communities in Lake Huron over a 5-year time period (2001–2005) in response to biotic and abiotic environmental factors. The association of environmental and spatial variables with trophic, life-history, and thermal traits at two spatial scales (regional basin-scale, local site-scale) was quantified using multivariate statistics and variation partitioning. We defined these two scales (regional, local) on which to measure variation and then applied this measurement framework identically in all 5 study years. With this framework, we found that there was no change in the spatial scales of fish community traits over the course of the study, although there were small inter-annual shifts in the importance of regional basin- and local site-scale variables in determining community trait composition (e.g., life-history, trophic, and thermal). The overriding effects of regional-scale variables may be related to inter-annual variation in average summer temperature. Additionally, drivers of fish community traits were highly variable among study years, with some years dominated by environmental variation and others dominated by spatially structured variation. The influence of spatial factors on trait composition was dynamic, which suggests that spatial patterns in fish communities over large landscapes are transient. Air temperature and vegetation were significant variables in most years, underscoring the importance of future climate change and shoreline development as drivers of fish community structure. Overall, a trait-based hierarchical framework may be a useful conservation tool, as it highlights the multi-scaled interactive effect of variables over a large landscape.     

Effects of natural hydrological variability on fish assemblages in small Mediterranean streams: Implications for ecological assessment

Small Mediterranean streams are shaped by predictable seasonal events of flooding and drying over an annual cycle, and present a strong inter and intra-annual variation in flow regime. Native fish assemblages in these streams are adapted to this natural environmental variability. The distinction of human-induced disturbances from the natural ones is thus a crucial step before assessing the ecological status of these streams. In this aim, the present study evaluates the effects of natural hydrological variability on fish assemblages from disturbed and least disturbed sites in small intermittent streams of south Portugal. Data were collected over the last two decades (1996–2011) in 14 sites located in the Guadiana and Sado river basins. High variability of fish assemblages was strongly dependent on human-induced disturbances, particularly nutrient/organic load and sediment load, and on natural hydrological variability. Natural hydrological variability can act jointly with anthropogenic disturbances, producing changes on fish assemblages structure of small intermittent streams. In least disturbed sites, despite the natural disturbances caused by inter-annual rainfall variations (including drought and flood events), fish assemblages maintained a long-term stability and revealed a high resilience. On the contrary, disturbed sites presented significantly higher variability on fish assemblages and a short and long-term instability, reflecting a decrease on the resistance and resilience of fish assemblages. Under these conditions, fish fauna integrity is particularly vulnerable and the ecological assessment may be influenced by natural hydrological variations. High hydrological variability (especially if it entails high frequency of dryer years and meaningful cumulative water deficit) may affect the impact of the human pressures with significant and consistent consequences on fish assemblage composition and integrity. In this study, fish metrics that maximize the detection of human degradation and minimize the response to natural variability were based on the relative abundance of native species (insectivorous species, eurytopic species, water column species, native lithophilic species), relative abundance of species with intermediate tolerance and relative number of exotic species. Results highlight the importance of assessing temporal variability on stream biomonitoring programs and emphasize the need to improve the assessment tools, accounting for long-term changes in fish assemblages, namely by selecting the most appropriate fish metrics that respond to anthropogenic disturbances but exhibit low natural temporal variability, essential both in the characterization of the biological reference conditions and in the development of fish indexes in intermittent streams.     

黄山陈村水库上游河源溪流的鱼类群落及其纵向梯度格局

确定鱼类群落的分布格局及其对人类活动的响应,是合理保护、恢复和管理鱼类多样性的基础。基于2011年5月和10月自黄山陈村水库上游3条河源溪流共39个样点的调查数据,比较研究了溪流间鱼类群落及其纵向梯度格局的异同,着重探讨了人类活动对溪流鱼类群落纵向梯度格局的影响。研究结果显示,同人为干扰较轻的舒溪相比,人为干扰严重的浦溪和麻溪中水宽、底质和植被覆盖率等局域栖息地条件显著变化,这造成了后者的鱼类多样性显著下降及物种组成的显著变化,主要表现为敏感性的地方物种(如宽鳍鱲、光唇鱼、原缨口鳅等)数量减少、耐受性的广布物种(如泥鳅、麦穗鱼、高体鰟鲏等)数量增多。舒溪的鱼类物种数及其组成均与海拔显著相关,但这种"海拔-鱼类群落"关系在麻溪和浦溪中削弱甚至消失。底质、植被覆盖率对舒溪鱼类群落具有重要影响,但对浦溪和麻溪鱼类群落却无显著影响。研究结果表明,在子流域空间尺度上,诸如城镇化发展、土地利用、河道治理等人类活动可通过对局域栖息地条件的影响,导致溪流鱼类多样性下降及其物种组成的变化,破坏鱼类群落的纵向梯度格局,并改变栖息地与鱼类群落之间的联系。     

Stream inflow and predation risk affect littoral habitat selection by benthic fish

1. We examined small, fishless headwater streams to determine whether transport of macroinvertebrates into the littoral zone of an oligotrophic lake augmented food availability for Cottus asper, an abundant predatory fish in our study system. We sampled fish and macroinvertebrates during the recruitment and growth season of 2 years, either monthly (2004) or bi‐monthly (2005), to observe whether stream inputs increased prey availability and whether variation in total macroinvertebrate biomass was tracked by fish. 2. Observations from eight headwater streams indicated that streams did not increase the total macroinvertebrate biomass in the shallow littoral zone at stream inflows, relative to adjacent plots without stream inputs (controls). The taxonomic composition of stream macroinvertebrates drifting toward the lake differed from that in the littoral lake benthos itself, although there was no evidence of any species change in the composition of the littoral benthos brought about by stream inputs. 3. Although streams made no measurable contribution to the biomass or taxonomic composition of the littoral macroinvertebrate benthos, there was substantial temporal variation in biomass among the eight sites for each of the (n = 7) sample periods during which observations were made. Variation in total biomass was primarily a function of bottom slope and benthic substrata in the lake habitats. Dominant taxonomic groups were Baetidae, Ephemerellidae (two genera), Leptophlebiidae, Chironomidae (three subfamilies) and Perlodidae, although we did not determine the specific substratum affinities of each taxon. 4. Mixed effects linear models identified a significant interaction between macroinvertebrate biomass and plot type (stream inflow vs. control) associated with fish abundance. Across the observed range of macroinvertebrate biomass, fish showed a significant preference for stream inflows, but more closely tracked food availability in the controls. For young‐of‐the‐year (YOY), a negative effect of temperature was also included in the model, and we observed lower temperatures at stream inflows. However, abundance of predatory adults affected habitat selection for YOY. Lake‐bottom slope also accounted for variation in abundance in both fish models. 5. Our results suggest that the effect of fishless headwater streams on downstream fish may not always be through direct delivery of food. In this study system, fish preferred stream inflow plots, but this preference interacted with macroinvertebrate biomass in a manner that was difficult to explain. For YOY, predation risk was related to the preference for stream inflows, although the specific factor that mitigates predation risk remains poorly understood.     

Spatiotemporal dynamics in a seasonal metacommunity structure is predictable: the case of floodplain‐fish communities

The metacommunity framework has greatly advanced our understanding about the importance of local and regional processes structuring ecological communities. However, information on how metacommunity structure and the relative strengths of their underlying mechanisms change through time is largely lacking. Dynamic systems that undergo environmental temporal changes and disturbances, such as floodplains, serve as natural laboratories to explore how their metacommunity structure change in time. Here we applied the Elements of Metacommunity Structure framework and variation partitioning analysis to assess how temporal changes in the local environmental factors and regional dispersal processes in the rain season influence a seasonal floodplain‐fish metacommunity. Across four months, relevant environmental factors were measured across 21 patches where over 3500 individual fish were sampled. Connectivity was measured using landscape resistance‐based metrics and additional spatial variation in metacommunity structure was assessed via spatial autocorrelation functions. The metacommunity structure changed from nestedness, at the beginning of the flood season, to a quasi‐Clementsian gradient at the end. Our analyses show that connectivity is only important in the beginning of the flood season whereas environment is only important at the end. These results suggest that this metacommunity is structured by changes between dispersal limitation and environmental filtering through time.     

The relative influence of local to regional drivers of variation in reef fishes

In this study, fishes and habitat attributes were quantified, four times over 1 year, on three reefs within four regions encompassing a c. 6° latitudinal gradient across south-western Australia. The variability observed was partitioned at these spatio-temporal scales in relation to reef fish variables and the influence of environmental drivers quantified at local scales, i.e. at the scale of reefs (the number of small and large topographic elements, the cover of kelp, fucalean and red algae, depth and wave exposure) and at the scale of regions (mean and maximum nutrient concentrations and mean seawater temperature) with regard to the total abundance, species density, species diversity and the multivariate structure of reef fishes. Variation in reef fish species density and diversity was significant at the regional scale, whereas variation in the total abundance and assemblage structure of fishes was also significant at local scales. Spatial variation was greater than temporal variation in all cases. A systematic and gradual species turnover in assemblage structure was observed between adjacent regions across the latitudinal gradient. The cover of red algae within larger patches of brown macroalgae (a biological attribute of the reef) and the number of large topographic elements (a structural attribute of the reef) were correlated with variation observed at local scales, while seawater temperature correlated with variation at the scale of regions. In conclusion, conservation efforts on reef fishes need to incorporate processes operating at regional scales with processes that shape local reef fish communities at local scales.     

Intraspecific life history variation in the threatened Louisiana pearlshell mussel, Margaritifera hembeli

1. Margaritifera hembeli is a threatened mussel limited to twenty-two headwater streams in the Red River drainage in central Louisiana, USA. This study evaluated intraspecific variation in density, growth, size and age structure and shell morphology among several isolated populations. This study also identified the host fish and considered the role that host fish distribution played in determining mussel recruitment. 2. Mussels were aggregated in beds and average densities differed among streams. However, maximum mussel densities in beds were similar in all streams; the observed maxima were among the largest for monospecific mussel beds in North America, often exceeding 300 individuals m^–2. 3. The maximum size reached by individuals differed among streams, but all size distributions were skewed towards larger individuals. A repeated measures analysis of tagged mussels in four populations, over a 2-year period, indicated 2-fold differences in growth rates among streams, and significant variation among years. Growth rates were not affected by local population density. Maximum ages reached, determined indirectly by comparing growth rates, varied from 45 to 75 years. A canonical discriminant analysis also revealed significant differences in shell morphology across populations. 4. Half of the populations showed evidence of recent recruitment, and these sites had fish assemblages dominated by the host fish Noturus phaeus (Taylor). Host fish abundance appeared more important than adult mussel density in explaining recruitment patterns. 5. Considerable intraspecific life history variation suggests that management strategies for this species should be stream-specific, with emphasis on ensuring long-term habitat stability.     

Testing the enemy release hypothesis: trematode parasites in the non-indigenous Manila clam <Emphasis Type="Italic">Ruditapes philippinarum</Emphasis>

Knowledge of temporal variation in nearshore Laurentian Great Lakes fish assemblages is important for understanding species–habitat associations, how abiotic and biotic influences vary temporally, and when sampling should occur. Using spring and fall seining data from Lake Erie beaches, we compared day and night fish assemblages and tested for differences among sampling periods. Beaches were utilized by a diverse collection of Lake Erie basin fishes (one-third of known species). During all sampling periods, catches were dominated by cyprinid species (53–91%), and by invertivores and planktivorous fishes. Diel differences were detected in abundance, species richness and assemblage structure. Multivariate analyses (canonical analysis of principal coordinates) indicated that season had a larger influence on fish assemblage structure than diel period. Given observed temporal variation in assemblage structure, studies of Laurentian Great Lakes beach fishes should be restricted to a single time period (e.g. day-time spring sampling), or adopt sampling designs that permit diel period and season to be included as factors in analyses. Second, the large seasonal variation in assemblage composition combined with higher night species richness indicates that night sampling during both spring and fall would be the most efficient and comprehensive approach for beach fish inventory. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Handling editor: J. Trexler     

Spatial and temporal variation of fish assemblage in two intermittent streams in north‐western Zimbabwe

This study examined the spatial and temporal variation in fish assemblage composition within two intermittent streams in north‐western Zimbabwe. Eight sites were sampled during three different periods of flow. Twenty‐five species in eight families, dominated by cichlids and cyprinids, were collected. Redundancy analysis (RDA) was used to explore the relationship between species and their environmental correlates. According to variation partition, 68.1% of assemblage variation was explained by both habitat and temporal variables. Most species were associated with large pools, whereas small species were associated with habitats that had gravel and boulders. The temporal aspect suggested a pattern associated with high catch per unit effort (CPUE) for the mormyrids, Clarias gariepinus and Labeo altivelis, during the early rain period. By comparison, the late rain period was associated with high CPUE for Labeo cylindricus, Labeobarbus marequensis, Chiloglanis neumanni and Oreochromis mortimeri, whereas the rest of the species had high CPUE during both the late rain and dry periods. The assemblage composition was more diverse during the late rain period. Refuge availability during the dry period appeared to be essential for the assemblage of the intermittent streams. Protection of these temporal habitats is therefore important in conserving fish assemblages within intermittent streams of the Lake Kariba area.     

Seasonal influence of brook trout and mottled sculpin on lower trophic levels in an Appalachian stream

1. In some situations fish have strong top‐down effects in stream communities while in others they seem to be relatively unimportant. Differences in the impact of fish may depend on a variety of factors including the foraging mode of the fish, interactions among fish species and temporal variation in environmental conditions and species interactions. 2. We investigated the effect of brook trout (Salvelinus fontinalis) and mottled sculpin (Cottus bairdi) on lower trophic levels in Appalachian streams and whether or not interactions between these fish changed their influence. Mesocosms were placed in a headwater stream in a randomized complete block design. Within blocks, mesocosms were randomly assigned to one of the following treatments: (i) no fish; (ii) sculpin only; (iii) trout only and (iv) both sculpin and trout. Fish biomass was the same in all three fish treatments. Invertebrate density and algal biomass in mesocosms were determined after 3 weeks. We repeated the experiment in the autumn, spring and summer to test for seasonality of fish effects. 3. The effect of fish on invertebrate assemblages was seasonal and depended on prey identity. Sculpin strongly suppressed grazer abundance in spring while trout had little effect on grazers in any season. The influence of both fish on insect predators was similar and relatively constant across seasons. We found little evidence of an interaction between sculpin and trout that strongly influenced their effect on prey across seasons. 4. None of the fish treatments influenced algal biomass during any of the seasons. Algal growth was also seasonal, with a two‐ to four‐fold increase in algal biomass in spring compared to autumn and summer. 5. Our results indicate that benthic and drift feeding fish differ in their effects on some, but not all prey. Furthermore, fish effects on prey were strongly seasonal for some, but not all prey types. While the temporal context is not commonly considered, our results indicate seasonality can be an important component of predator–prey interactions in streams.     

Scales of spatiotemporal variability in macroinvertebrate abundance and diversity in monsoonal streams: detecting environmental change

1. We quantified spatial and temporal variability in benthic macroinvertebrate species richness, diversity and abundance in six unpolluted streams in monsoonal Hong Kong at different scales using a nested sampling design. The spatial scales were regions, stream sites and stream sections within sites; temporal scales were years (1997–99), seasons (dry versus wet seasons) and days within seasons. 2. Spatiotemporal variability in total abundance and species richness was greater during the wet season, especially at small scales, and tended to obscure site‐ and region‐scale differences, which were more conspicuous during the dry season. Total abundance and richness were greater in the dry season, reflecting the effects of spate‐induced disturbance during the wet season. Species diversity showed little variation at the seasonal scale, but variability at the site scale was apparent during both seasons. 3. Despite marked variations in monsoonal rainfall, inter‐year differences in macroinvertebrate richness and abundance at the site scale during the wet season were minor. Inter‐year differences were only evident during the dry season when streams were at base flow and biotic interactions may structure assemblages. 4. Small‐scale patchiness within riffles was the dominant spatial scale of variation in macroinvertebrate richness, total abundance and densities of common species, although site or region was important for some species. The proportion of total variance contributed by small‐scale spatial variability increased during the dry season, whereas temporal variability associated with days was greater during the wet season. 5. The observed patterns of spatiotemporal variation have implications for detection of environmental change or biomonitoring using macroinvertebrate indicators in streams in monsoonal regions. Sampling should be confined to the dry season or, in cases where more resources are available, make use of data from both dry and wet seasons. Sampling in more than one dry season is required to avoid the potentially confounding effects of inter‐year variation, although variability at that scale was relatively small.     

Scales of spatiotemporal variation in macroinvertebrate assemblage structure in monsoonal streams: the importance of season

1. We assessed the relative importance of different scales of spatial and temporal variability on benthic macroinvertebrate assemblage structure in six unpolluted streams in monsoonal Hong Kong using ordination and complementary multivariate analyses. The spatial scales were regions, sites (streams) and sections (riffles) within sites. The temporal scales were years (three, including one with unusually high rainfall), seasons (dry versus wet) and days within seasons. 2. Significant differences in assemblage structure were manifested at all temporal scales. Those at the site scale were most obvious, whereas demarcation of assemblage structure at the section (riffle) scale was smaller, and there was no significant regional differentiation in assemblage structure. Seasonal variability in assemblage structure was greater than that among years or days. 3. Inter‐year differences in assemblage structure were recorded at all sites, and were noted among all years at some sites but not at others. They were recorded more frequently during the dry season, although their occurrence (in pair‐wise comparisons between years) appeared to be related to differences in the monsoonal (wet season) rainfall. 4. Seasonal differences in assemblage structure were strongly evident at all sites. Inter‐site differences were more apparent during the dry season when local (site‐scale) influences on assemblages were stronger. By contrast, wet‐season samples were more variable because of spate‐induced disturbance, and inter‐site differentiation was less distinct. 5. Differences among days at all sites were relatively minor, but shifts attributable to repeated spate‐induced disturbance were evident at some sites during the wet season. 6. Differences at the section scale were recorded more frequently during the dry season, when the extent of within‐site variability among sections was higher, reflecting increased patchiness within sections resulting from increased substratum heterogeneity and/or greater intensity of biotic interactions. 7. Seasonal shifts in macroinvertebrate assemblage structure at a variety of scales in Hong Kong streams are likely to be attributable to monsoonal rains affecting the relative intensity of abiotic disturbance and biotic interactions in accordance with the harsh‐benign hypothesis.