The influence of abiotic factors and density-dependent mechanisms on between-year variations in a stream invertebrate community

Samples of invertebrates were taken in two seasons (October and April) over 5 years in a trout stream in Belgium. Diversity, total benthic invertebrate density, and densities of 27 of 34 taxa tested showed significant differences between years. Significant seasonal effects and interactions were observed in 18 and 7 taxa respectively. The most common pattern of between-year variation was related to hydrological events (flooding) which caused low densities for many species. Most species recovered rapidly, but the recovery of some predators seemed to show a delay of one year. The amphipod Gammarus fossarum appeared to respond positively to flooding as well as to certain post-flood conditions. A significant positive correlation between mortality and the logarithm of initial density was obtained for 6 of 11 species tested. The relationships between mortality and abiotic factors such as water temperature, mean and maximum water level, were very poor. A hierarchy of environmental factors in the regulation of stream macroinvertebrate populations is suggested with: (1) exceptional abiotic events acting in a density-independent manner; and (2) biotic factors that may account for density-dependent regulation.     

Invertebrate Sampling in the Substratum of an Experimental Recirculating Stream

In order to determine the efficacy of sampling benthic invertebrates to a depth of 100 mm as compared with 200 mm, the gravel substratum of an experimental recirculating stream was sampled to both depths. Except for 6 taxa which were significantly more numerous in the shallower samplers, there was no significant difference between the densities determined from the two depths.     

Effects of experimental flow regulation on invertebrate drift and stranding in the Flathead and Kootenai Rivers,Montana, USA

Studies were conducted to determine the effects of experimental manipulations of discharge on invertebrate drift in two regulated rivers in northwestern Montana, USA. During these studies the discharge regime in the Flathead River was characterized by frequent flow fluctuations, while in the Kootenai River high discharge was maintained for much longer periods before flow was reduced to minimum discharge. The magnitude of the response of invertebrates to disturbance was different in the two rivers, in part because of the different frequencies of flow changes. Midstream invertebrate drift increased an order of magnitude during increasing discharges in the Flathead River but was not substantially increased during decreasing discharges. When the prior discharge regime had been sustained at high levels in the Kootenai River, invertebrate drift densities as high as 300 000/100 m³ were measured along the shoreline following reductions in discharge, both immediately after flow began to decrease and after dark on the same day. There was also more recolonization of shoreline areas and more stranding of insects following dewatering of nearshore regions when there had been sustained high discharge levels prior to the flow reduction. More insect stranding occurred during a faster rate of decrease in discharge (50 000 to 100 000 organisms m⁻²).     

Habitat structure and invertebrate assemblages on stream stones: A multivariate view from the riffle

Abstract Highly structured habitats, those with complex or heterogeneous physical structure, commonly contain more species than simply structured ones. However, tests of this relationship have been hampered by habitat-specific definitions and measurements of habitat structure and, in stream studies, by insufficient information on spatial variation in faunal abundances. We sampled the fauna of 90 stones collected from an order 4 section of the Taggerty and Steavenson Rivers, southeastern Australia. The spatially nested sampling design encompassed three spatial scales (sites within the same stream order, riffles at the same site and groups of stones within the same riffle) but no temporal replication, as we were only able to sample once. Habitat structure of stones was quantified by measuring the amount and type of epilithon on stones, together with stone shapes, sizes, textures and surface complexities. For the latter two attributes, we used a new method, involving image analysis, which can be used to quantify habitat structure in comparable ways in other systems. Species richness (S) was related to some measures of habitat structure, being higher on stones with a rough texture and weakly related to the amount of epilithon present. Total numbers of individuals (N) were also higher on rough stones, and faunal composition (as quantified by hybrid multidimensional scaling) was related to abundance of epilithon. However, flow environments, as quantified by water velocities and depths, were also related to faunal composition. Riffles varied in both faunal densities and near-bed flow environment, but not in any consistent way. Such variability means that individual riffles cannot be reasonably used to ‘represent’ stream sections or orders, an assumption commonly made in stream studies. Neglect of such issues has also resulted in poor conceptual integration between large- and small-scale studies in stream ecology.     

A continuous study of the total drift of freshwater shrimps, Gammarus pulex, in a small stony stream in the English Lake District

1. The objective was to determine the major factors affecting the downstream dispersal (drift) of freshwater shrimps, Gammarus pulex. Sample replication and frequency are major problems in the quantification of drift. For the first time, these problems were avoided by sampling the whole stream continuously so that all the shrimps drifting downstream at the sampling point were caught in a net emptied at dusk and dawn in 1966, and every 3 days in 1967. 2. There was no consistent seasonal pattern in drift rates, but a high proportion of annual drift was taken in only a few samples. There was a nocturnal diel pattern of drift with peaks soon after dusk and just before dawn. A power function described the significant (P < 0.001) relationship between drift and flow, and was used to neutralise the dominant effects of flow by standardising total drift over 24 h, nocturnal drift and diurnal drift (drift per 50 m³). These were all significantly (P < 0.001) related to benthos density, but not to date, temperature, or length of the night or day. 3. The relationship between drift values and the independent variables, flow and benthos density, was well described (P < 0.001) by a multiple‐regression model. Adding temperature, date, and/or the length of the night or day did not improve model fit. Variations in flow and benthos density explained 94% of the variation in total drift over 24 h, 97% of the variation in nocturnal drift, but only 44% of the variation in diurnal drift. A power function described (P < 0.001) the relationship between total drift and the volume of water sampled over 3‐day periods in 1967. Flow explained 95% of this drift variation; it was unnecessary to add another variable such as benthos density. 4. The significance of this study is that it avoided the problems associated with the quantification of drift samples. Therefore, the conclusions are more robust than those of many previous studies. A high proportion of the annual drift losses would have been undetected by intermittent sampling. Temperature, season, night or day length had no significant effect on drift densities, and the relationship between drift and benthos densities was proportional, not density dependent. The nocturnal increase in drift could not be interpreted as an antipredator behaviour. The dominance of flow and benthos density was apparent but the quantitative relationships posed further questions, especially those related to drift distances at different velocities.     

A study on the faeces of some chalk stream invertebrates

An account is given of the size, form, texture, colour, cohesion and composition of the faeces of 41 species of invertebrates found in chalk streams. The relationships between the character of the faeces and the taxonomic positions and habits of the producers are considered.The importance of faeces production relative to the bulk of stream bed sediments is discussed. Faeces of different origins accumulate in different areas of the stream bed and these differences may be associated in part with their form and structure and in part with the distribution of the species from which they originate. In the summer months tubificid worms alone may be responsible for reworking between 0.3% and 0.5% of the fine particulate material in sediments every day.     

The influence of hydrology on the structure of invertebrate communities in two streams flowing into Lake Nakuru,Kenya

Stream fauna communities were studied in the Baharini Springbrook and the Njoro River, which flow into Lake Nakuru. The streams have significantly different flow regimes. Consequently, discharge-related factors influence their faunal composition and abundance. The Baharini Springbrook is perennial, slow-flowing with low steady discharge, a higher conductivity and a low conductivity gradient along its course. The Njoro River is intermittent, fast-flowing with a higher and variable discharge, lower conductivity and a steeper conductivity gradient. Due to its permanent and steady surface stream flow, the Baharini Springbrook has a higher number of invertebrate species and higher abundances than the Njoro River. However, in spite of the intermittent nature of the Njoro River, during periods of relatively steady discharge the composition of its fauna is similar to that of the Baharini Springbrook. Similarity in the substratum characteristics of the two streams and their close proximity is a possible explanation for the close relationship in their faunal composition. At the onset of flow after a drought, rapid colonisation of the lowland intermittent stretch of the Njoro River occurred probably by drift from upstream perennial sections and/or from the hyporheos. Initial small spates at the onset of surface flow seem not to affect the invertebrate abundance, but later bigger spates during and after much heavier rains reduced the faunal abundance significantly. The recovery of faunal abundance to pre-flood levels in the Njoro River was completed in 27 days. Taxon richness was, however, less affected by the spates.     

Land‐use legacy and the differential response of stream macroinvertebrates to multiple stressors studied using in situ experimental mesocosms

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An analysis of changes in the invertebrate community along a southern New Zealand montane stream

Changes in benthic community structure at four sites along a flood-prone montane stream continuum in the South Island of New Zealand were analysed using rank correlation (Kendall's Tau and Spearman's r_s) and a community similarity index (S_c) developed initially for stream pollution assessment. Species richness was highest in a forested headwater stream, which is attributed to its relatively greater physical stability and habitat heterogeneity. Community similarity decreased with distance apart, and pairs of open and forested sites showed the greatest similarity. Species composition and relative abundance along the continuum changed least in the numerically dominant Ephemeroptera and Plecoptera, groups in which several species had broad habitat requirements. In contrast, a number of dipteran and trichopteran species had more restricted distributions. The more specialised niche requirements of some species in these orders are attributed to their generally small size, their ability to spin silk, and their holometabolous development. Physical rather than biological factors appeared to limit the distributions of most invertebrates, and there was little evidence of ecological segregation by either food resource partitioning or differences in life history patterns amongst many closely related species. Theories that stream communities are well ordered entities maximising the efficiency of energy utilisation cannot realistically be applied to such unstable stream systems.     

Comparison of stream benthic invertebrate assemblages among forest types in the temperate region of Japan

We compared benthic invertebrate assemblages among headwater streams in several forest types in Japan. Forests were divided into three clusters based on vegetation composition: old-growth broad-leaved forest, planted coniferous forest, and mixed forest. The numbers of individuals and families and the diversity (Shannon-Wiener) of benthic invertebrate assemblages did not differ significantly among the three forest clusters. However, principal components analysis of family abundance showed differences in the benthic invertebrate assemblages among the three forest clusters. No environmental factors were correlated with these differences. Benthic invertebrate assemblages differed depending on forest composition. The abundances of Taeniopterygidae and Athericidae in old-growth broad-leaved forest were significantly greater than in planted coniferous forest. The abundances of Heptageniidae, Baetiidae, Stenopsychidae, Uenoidae, Chironomidae, and Potamidae in planted coniferous forest were significantly greater than in old-growth broad-leaved forest. If the remaining old-growth broad-leaved forest were to be converted to coniferous plantation, species that inhabit old-growth forest may become extinct.     

Relative roles of stream flow and sedimentary conditions in controlling hyporheic exchange

Hyporheic exchange is often controlled by subsurface advection driven by the interaction of the stream with sedimentary pore water. The nature and magnitude of the induced exchange flow is dependent on the characteristics of both the stream flow and the sediment bed. Fundamental hydrodynamic theory can be applied to determine general relationships between stream characteristics, sediment characteristics, and hyporheic exchange rates. When the stream bed is fine enough to allow application of Darcy's Law, as with sand beds, the induced advective exchange can be calculated from fundamental hydrodynamic principles. Comparison with a wide range of experimental results demonstrates the predictive capability of this theory. Coarser sediments such as gravels are more complex because they admit turbulent interactions between the stream and subsurface flows, which can produce considerable exchange even when the bed surface is flat and no flows are induced by the bed topography. Even for this case, however, scaling arguments can still be used to determine how exchange rates vary with stream and sedimentary conditions. Evaluation of laboratory flume experiments for a wide range of stream conditions, bed sediment types including sand and gravel, and bed geometries demonstrates that exchange scales with the permeability of the bed sediments and the square of the stream velocity. These relationships occur due to fundamental hydrodynamic processes, and were observed to hold over almost five orders of magnitude of exchange flux. Such scaling relationships are very useful in practice because they can be used to extend observed hyporheic exchange rates to different flow conditions and to uniquely identify the role of sedimentary conditions in controlling exchange flux.     

Temporal changes in benthic macroinvertebrates and their interactions with fish predators after restoration in the Cheonggyecheon,a downtown stream in Seoul,Korea

The Cheonggyecheon (Cheonggye stream), a downtown stream in Seoul, Korea, was restored by the historic Cheonggyecheon Restoration Project (2003–2005), and its discharge and other environmental conditions are artificially regulated throughout the year. The aim of this study was to assess temporal changes in benthic macroinvertebrates and their interactions with fish predators in the Cheonggyecheon. Sampling was conducted twice a year (spring and autumn) at three sections of the stream from 2006 to 2010 using a Surber sampler (50 × 50 cm) for benthic macroinvertebrates and a cast net (mesh size, 10 × 10 mm) and a kick net (mesh size, 5 × 5 mm) for fishes. Analyses of annual precipitation and most water quality parameters showed no significant differences across the study period (P > 0.05). The species richness and density of benthic macroinvertebrates and fishes varied across the sampling period. Analyses of functional feeding groups and trophic guilds showed distinct trends, such as an increase in the number of insectivorous fish in the absence of invertebrate predators. Correlation and regression analyses for five major groups of benthic macroinvertebrates and fishes demonstrated that the density of certain groups of benthic macroinvertebrates (e.g. Tubificidae, Erpobdellidae, Baetidae, and Chironomidae) and insectivorous fishes (e.g. Gobioninae, Leuciscinae, Danioninae, and Gobiidae) were negatively correlated. In conclusion, in an artificially regulated stream like the restored Cheonggyecheon, where physical environments are nearly unchanged, fish predation can be a major factor controlling benthic macroinvertebrate communities.     

Seasonal aspects of the invertebrate fauna of three species of plants and rock surfaces in a small stream

Macroinverbrates associated with three species of aquatic plants and those on rock surfaces in a stream were collected every two weeks from late May to late November. Some animals appeared to use all plants opportunistically as substrates from which food sources could be better exploited throughout the season. Others, usually of minor numerical importance, strongly differentiated between plant species and appeared to find the resources that they may require on only one plant and only at specific times of the year. Myriophyllum alterniflorum supported the highest density of invertebrates but numbers per unit surface area fluctuated widely throughout the season. Numbers of individuals on exposed rocks were remarkably constant and animals that dominated there were usually different from those on plants. Patterns in both total individuals and numbers of individuals within different taxa indicate that seasonal variation in phytomacrofauna in streams is significant and should be considered in the design of future studies.     

Comparative tests of the species‐genetic diversity correlation at neutral and nonneutral loci in four species of stream insect

A fundamental question linking population genetics and community ecology is how adaptive processes (e.g., natural selection) and neutral processes (e.g., drift‐migration equilibrium) underpin the species‐genetic diversity correlation (SGDC). Here, we combine genome scans and outlier loci detection with community analysis to separately test for neutral and nonneutral SGDCs in four species of stream insect. We sampled 60 localities in Japan and examined the relationships among population AFLP band richness (Br), taxon richness of the total community (S) and of the trophic guild (S_tr), and 15 habitat parameters that could potentially drive adaptation and influence richness. Neutral Br was positively correlated with S only in the dominant species of these communities, suggesting Br may be constrained when intraspecific competition is pronounced. Nonneutral Br was correlated with S_tr in a species restricted to high elevations where habitat heterogeneity was highest. Community distance and genetic distance (β‐SGDC) was correlated in two of the four species at both neutral and nonneutral loci. Distance‐based redundancy analysis found geographic isolation and elevation to drive divergence of both communities and populations. This suggests that both neutral and adaptive divergence occurred through the shared influences of geographic isolation and local adaptation at the two levels of diversity.     

长江口及其邻近海域春季无脊椎动物群落时空变化

为了研究长江口及其邻近海域无脊椎动物群落结构和多样性的时空变化特征, 我们于1999-2012年春季在长江口及其邻近海域采用定点底层双拖网调查方式进行无脊椎动物调查。结果表明: (1)1999-2012年长江口水域共记录无脊椎动物41种, 隶属6纲10目23科, 其中甲壳动物种类最多(26种), 其次为软体动物(13种)。不同年份种类数量呈先下降后上升的趋势。(2)优势种主要包括日本枪乌贼(Loligo japonica)、三疣梭子蟹(Portunus trituberculatus)、葛氏长臂虾(Palaemon gravieri)和鹰爪虾(Trachypenaeus curvirostris)等, 其在年间存在剧烈变动, 但日本枪乌贼几乎每年都是优势种。(3)长江口无脊椎动物丰度、种类丰富度和多样性在年间均存在显著差异, 1999年和2001年最高, 2004年后呈先下降后恢复上升的趋势。(4)长江口水域的无脊椎动物在每个航次调查中都存在2-3个群聚类型, 并有不同的指示种类。(5)1999-2012年长江口无脊椎动物群落的时间变化可划分为3个阶段: 1999-2001年多样性程度最高, 2004-2007年下降至最低水平, 2009-2012年多样性显著回升, 但尚未恢复到1999-2001年的水平。与20世纪80年代相比, 蟹类减少导致长江口无脊椎动物生物量整体水平下降, 高营养级生物资源衰退带来了无脊椎动物中低营养级生物种群的迅速发展。     

Size‐related and diel variations in microhabitat use of three endangered small fishes in a Mediterranean coastal stream

This study analysed the microhabitat use of three endangered fish species, Andalusian toothcarp Aphanius baeticus , Iberian loach Cobitis paludica and sand smelt Atherina boyeri , in a coastal stream stretch. Plastic minnow traps were set both during daytime and at night on the bottom and at the surface. Depth and presence of effective refuge were recorded for each trap. To assess size‐related changes in microhabitat use individuals of each species were classified in three size classes. The three species preferentially used bottom positions in the water column, though this behaviour was more evident in the case of Iberian loach. While large Iberian loach remained active at night Andalusian toothcarp and sand smelt were strongly diurnal, especially larger individuals. The three species showed a clear ontogenetic change in microhabitat preferences towards deeper waters. Small Andalusian toothcarp and medium‐sized Iberian loach used deeper microhabitat in the presence of refuge. Large Andalusian toothcarp consistently preferred exposed microhabitat at any time. Andalusian toothcarp using refuge were smaller at any time, while Iberian loach followed this pattern only at night. The differential vulnerability of these species to different predators (aerial and aquatic; diurnal and nocturnal) could explain the observed patterns in microhabitat use. Fish tended to co‐occur in microhabitats either due to habitat characteristics independently of species or due to species independently of habitat. Andalusian toothcarp segregated spatio‐temporally from sand smelt and Iberian loach, but these species occurred independently of each other. According to these co‐occurrence patterns, Andalusian toothcarp would be more sensitive than Iberian loach or sand smelt to interspecific interactions.     

Drought intensification alters the composition,body size,and trophic structure of invertebrate assemblages in a stream mesocosm experiment

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Day–night changes in the spatial distribution and habitat preferences of freshwater shrimps, Gammarus pulex, in a stony stream

1. As many invertebrates are nocturnal, their spatial distribution and habitat preferences may change from day to night. Both aspects are examined for Gammarus pulex by testing the hypotheses: (i) a power function was a suitable model for the spatial distribution of the shrimps in both day and night; (ii) diurnal and nocturnal spatial distributions were significantly different; (iii) diurnal and nocturnal habitat preferences were significantly different. Five different life‐stages were treated separately. To ensure that the conclusions were consistent, large samples were taken near midday and midnight in April, June and November over 4 years at two sites about 3 km apart in a stony stream: downstream (n = 30) and upstream (n = 50). 2. The first and second hypotheses were supported at both sites. A power function, relating spatial variance (s²) to mean (m), was an excellent fit in all analyses (P < 0.001, r² > 0.91), i.e. the spatial variance was density‐dependent. All five life‐stages were aggregated in the day. At night, the degree of aggregation increased for juveniles at higher densities but decreased for juveniles at lower densities, increased for immature females and males, but decreased slightly for mature females and especially mature males, the latter being close to a random distribution. There were no significant differences between sites, in spite of the lower numbers at the downstream site. 3. The third hypothesis was tested at only the upstream site and supported by comparisons between shrimp densities and 13 physical variables (distance from bank, water depth, water velocity, ten particle size‐classes), and three non‐physical variables (dry weights of bryophytes, leaf material, organic detritus). During the day, densities were strongly related to particle sizes with the following preferences: 0.5–8 mm for juveniles, 8–256 mm for the other life‐stages with a weaker relationship for males. There were no significant positive relationships with the other variables, apart from bryophytes for immature shrimps and adults. At night, densities were unrelated to particle size; juveniles and immature shrimps preferred low water velocities near the banks, often where leaf material and organic detritus accumulated, females often preferred medium water velocities slightly away from the banks, and males showed no habitat preferences. 4. Day samples do not provide a complete picture of habitat preferences and probably identify refuge habitats. Day–night changes in spatial distribution and habitat preferences are an essential part of the behavioural dynamics of the shrimps and should be investigated in other species.     

Drought ecohydrology alters the structure and function of benthic invertebrate communities in mountain streams

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Effects of pulsed and pressed disturbances on the benthic invertebrate community following a coal spill in a small stream in northeastern USA

In September 1999, a coal-carrying train derailed and spilled 180,000–270,000 kg of coal into the Cayuga Inlet near Ithaca, New York. This study determined the immediate effects of the coal spill and the clean up procedures on the aquatic invertebrate community, and whether the stream recovered from this event after 2 years. Benthic invertebrate samples were taken both upstream and downstream of the coal spill immediately following clean-up efforts and two years later. Just after the coal spill, the total abundance and species richness of aquatic invertebrates were significantly lower downstream of the spill, suggesting that the disturbance caused increased mortality and/or emigration compared to a reference site upstream. Taxa affected most were grazers and turbidity-susceptible invertebrates. Two years later the invertebrate communities were similar upstream and downstream of the spill, except for an increase in the percent of the dominant genus, Hydropsyche(Trichoptera: Hydropsychidae). We speculate that long-term effects of channelization of the stream that occurred during the clean-up prevented the invertebrate assemblage from returning to the conditions observed in a reference site upstream of the coal spill. We propose that large scale environmental clean-ups should be designed to avoid altering ecosystems permanently, and that streams should be allowed to recover naturally without destructive human intervention.