Showing papers in "Freshwater Biology in 1996"

A user‐friendly guide to the ciliates (Protozoa, Ciliophora) commonly used by hydrobiologists as bioindicators in rivers, lakes, and waste waters, with notes on their ecology

Abstract: SUMMARY 1A user-friendly guide to 300 ciliate species (Protozoa, Ciliophora) used as bioindicators by river, lake and waste water ecologists is provided. The guide is an English translation of the flow charts written in German and published by Foissner et al. (1991, 1992, 1994, 1995) in the Ciliate Atlas, a monograph on the ciliates used as bioindicators in the saprobic system. This guide is designed for users not specifically trained in identification of ciliates. Main groups and species are keyed dichotomously on forty-seven flow charts using simple characters usually recognizable in live specimens. Species with conspicuous characters, e.g. large size or distinct colour, are shown on thirty-two separate charts designated ‘special keys’. Although the flow charts give a high probability of correct species identifications, these should nevertheless be checked against the detailed figures and descriptions contained in the Ciliate Atlas. 2A table with the species keyed and their main ecological characteristics (biomass, food preference, salinity tolerance, preferred occurrence, saprobiological classification) is also provided. 3Typical ciliate communities found in natural and polluted habitats are briefly described and figured on thirteen plates. 4A detailed systematic index is provided for all taxa mentioned in the flow charts.

A hydrogeography of unregulated streams in the United States and an examination of scale‐dependence in some hydrological descriptors

Abstract: 1. Relatively undisturbed streams in continental U.S.A. were classified according to variation in ten ecologically relevant hydrological characteristics. Measures of flow variability and predictability for average conditions, as well as for low- and high-flow extremes, were extracted from long-term (15–58-year) daily streamflow data for 806 streams. 2. Two groups of sites were analysed: all 806 sites and a subset of 420 ‘best’ sites. For each group, cluster analysis identified ten distinctive stream types, seven permanent and three intermittent. The geographical clustering exhibited by the stream types indicated regional differences in climatic and geological features. A bootstrapping technique applied to the permanent stream classes showed the majority of them were statistically robust. 3. The derived classification of U.S. streams based on ecologically relevant hydrological characteristics provides a comprehensive catalogue of small to mid-size streams that, according to ecological theory, may differ in major aspects of their ecological organization. The classification provides a basis for hypothesis generation and affords an objective framework for matching streams for purposes of comparative ecological investigations. 4. A subset of 118 streams from the ten classification groups was selected to determine whether certain hydrological variables often used by ecologists to make cross-system comparisons are sensitive to the temporal coarseness of the hydrological time series used to derive the variables. The three hydrological variables considered were streamflow predictability, streamflow variability and flood timing. 5. Streamflow predictability (using Colwell’s Index) was calculated at daily, weekly, monthly and seasonal time scales. Estimates of predictability showed either no change across time scales, a gradual and consistent increase across time scales, or a maximum value at the monthly time scale. Coefficient of variation of streamflow was calculated at daily, weekly, monthly, seasonal, and annual time steps. Daily values were always greatest for all streams. Some groups showed minimum variability at the monthly scale, others at the seasonal. Timing of daily peak flows could be detected with 50–90% accuracy across stream groups using coarser grain monthly and annual hydrographic data. 6. Inferences about hydrological similarity among streams across broad geographical scales can be sensitive to choice of time scale used in the hydrological characterization.

Diel, episodic and seasonal changes in pH and concentrations of inorganic carbon in a productive lake

Abstract: SUMMARY 1 Two pH electrodes and a thermistor were used to record conditions in the surface of Esthwaite Water every 15 min over a 12-month period. Combined with approximately weekly measurements of alkalinity they allowed inorganic carbon speciation to be calculated. 2 Large changes in pH from 7.1 to nearly 10.3, and hence in concentrations of inorganic carbon species, were measured over a year. Carbon speciation and pH varied on a diel, episodic and seasonal basis. Diel variation of up to pH 1.8 was recorded, although typical daily variation was between 0.03 and 1.06 (5 and 95 percentiles). Daily change in concentration of inorganic carbon varied between 4 and 63 mmol m-3 (5 and 95 percentiles). 3 During lake stratification, episodes of high pH, typically of 1–2 weeks' duration were interspersed with episodes of lower pH. These changes appeared to relate to the weather: e.g. low wind velocity, high pressure, low rainfall and high sunshine hours correlated with periods of high pH. 4 Seasonal progression of carbon depletion generally followed stratification and the development of high phytoplankton biomass. When the lake was isothermal, the phytoplankton biomass caused relatively small amounts of carbon depletion. 5 During autumn, winter and spring, the lake had concentrations of CO2* (free CO2) up to 0.12 mol m-3 which is nearly seven times the calculated atmospheric equilibrium concentration so the lake will accordingly be losing carbon to the atmosphere. In contrast, during periods of elevated pH the concentration of CO2* was reduced close to zero and the lake will take up atmospheric CO2. The rates of transfer between water and the atmosphere were estimated using a chemical equilibrium model with three boundary layer thicknesses. The calculations show that over a year the lake loses CO2 to the atmosphere with the current mean atmospheric level of 360 μmol mol-1, at between 0.28 and 2.80 mol m-2 yr-1. During elevated pH, rates of CO2-influx increased up to nearly tenfold as a result of chemical-enhancement by parallel flux of HCO-3. Input of CO2* to the lake from the catchment is suggested to be the main source of the carbon lost to the atmosphere. 6 The turnover time for CO2 between the air and water was calculated to be 1 year for the gross influx and 3.3 years for the net flux. These values are less than the average water residence time of 0.25 years, which indicates that over a year inflow from streams is a more important source of inorganic carbon than the atmosphere. 7 Influx of CO2 from the atmosphere was calculated to be roughly equivalent to between 1 and 4% of the rates of production in the water during mid-summer indicating that this source of inorganic carbon is not a major one in this lake. 8 Influx of CO2 from the hypolimnion was estimated on one occasion to be 6.9 10-9 mol m-2 s-1 using transfer values based on mass eddy-diffusion. These rates are equivalent to 23% of the rate of influx of CO2 from the atmosphere on this occasion which suggests that the hypolimnion is probably a small source of inorganic carbon to the epilimnion. The exception appears to be during windy episodes when pH is depressed. Calculations based on depth-profiles of CO2* and HCO-3 suggest that the measured changes in pH can be accounted for by entrainment of hypolimnetic water into the epilimnion. 9 The solubility product for calcite was exceeded by up to about sixfold which, although insufficient to allow homogeneous precipitation, may have allowed heterogeneous precipitation around algal particles.

Patterns in benthic food webs: a role for omnivorous crayfish?

Abstract: 1. The biomass and species richness of macrophytes and invertebrates in artificial ponds at two sites in southern Sweden (twenty-one ponds at each site) were investigated. Alkalinity was high at one site (H ponds) and low at the other site (L ponds). The ponds chosen had different densities of signal crayfish (Pacifastacus leniusculus), with mean crayfish abundance (estimated by trapping and expressed as catch per unit effort) significantly higher in the L ponds (10.7) than in the H ponds (4.9). Macrophytes, invertebrates, the amount of periphyton on stones and the organic content of the sediment were determined in each pond. 2. Macrophyte biomass, cover and species richness declined with increasing crayfish density. Macrophyte species composition differed between ponds and was related to crayfish abundance. 3. The total biomass of invertebrates and the biomass of herbivorous/detritivorous invertebrates declined with increasing crayfish abundance, but the biomass of predatory invertebrates declined only in the L ponds. The relative biomass of Gastropoda and Odonata declined in ponds where crayfish were abundant. In ponds where crayfish were abundant the invertebrate fauna was dominated by sediment-dwelling taxa (Sialis (H and L ponds) and Chironomidae (H ponds)). 4. The number of invertebrate taxa in macrophytes declined with increasing crayfish abundance. The percentage of macrophyte-associated invertebrate taxa differed between ponds, but also between sites. The relative biomass of Gastropoda declined in H ponds where crayfish were abundant. In H ponds Trichoptera or Gammarus sp. and Heteroptera dominated where crayfish were abundant, whereas Odonata dominated in L ponds with abundant crayfish. 5. The organic content of the sediment decreased in ponds with high crayfish densities, while the amount of periphyton on stones was not related to crayfish density. 6. We conclude that the signal crayfish may play an important role as a keystone consumer in pond ecosystems, but lower trophic levels did not respond to changes in the abundance of the crayfish according to the trophic cascade model. Omnivorous crayfish may decouple the cascading effect.

The effect of habitat‐specific sampling on biological assessment of water quality using a predictive model

Abstract: 1. Detection of impairment in macroinvertebrate communities using rapid biological assessment depends on the ability to compare sites, with confidence that differences obtained result from water quality. However, collections from more than one habitat type may introduce variation that can potentially mask water quality differences among sites. Data were collected from the riffle, edge, pool-rock and macrophyte habitats at reference (minimally disturbed) and test (disturbed) stream sites throughout the Australian Capital Territory. The effect of habitat-specific sampling on predictive models for detecting impairment in macroinvertebrate communities was determined. Four models were used: riffle only, edge only, each habitat as an individual object, and all habitats sampled at a site considered as a composite sample. 2. Macroinvertebrates from individual habitats generally clustered into separate groups because collections from the same habitat at different sites were more similar than collections from different habitats within a site. Thus, in the habitats as individual objects model, the taxa predicted to occur at a test site may be an indication of habitat type rather than water quality. The outputs of the composite habitats and riffle and edge models were similar. However, the variable number of habitats included at each site in the composite model may confound the detection of biological impairment because of unequal sampling effort. The riffle and edge models were the most robust because they were less confounded by inter-habitat variation and were based on comparisons made between equivalent environmental units. 3. Comparison of observed/expected taxa ratios for test sites showed that each model could detect biological impairment, indicating considerable data redundancy was introduced by sampling several habitats. In particular, the pool-rock and macrophyte habitats contributed no information with regard to macroinvertebrate taxon occurrence or detection of biological impairment that could not be obtained from either the riffle or edge habitats within the study area.

Artificial mixing prevents nuisance blooms of the cyanobacterium Microcystis in Lake Nieuwe Meer, the Netherlands

Abstract: 1. Artificial mixing in the hypertrophic Lake Nieuwe Meer was successful in preventing blooms of the cyanobacterium Microcystis. During the 2 years of artificial, deep mixing the number of colonies of Microcystis per litre and also per m(2) was lower than in the two preceding control years. Hardly any nuisance scums of Microcystis occurred in the lake. 2. The phytoplankton shifted from a cyanobacteria-dominated community in summer to a mixed community of flagellates, green algae and diatoms. Reduced sedimentation losses in the mixed lake, probably in combination with a lower pH, favoured nonbuoyant algae, while the entrainment of cyanobacteria in the turbulent flow nullified their advantage of buoyancy. 3. The chlorophyll concentrations were much lower in the mixed lake, but the euphotic depth did not show clear differences between the years. The chlorophyll content integrated through depth (m(-2)) increased in the artificially mixed lake. 4. The deep lake normally stratified in summer, but artificial mixing of the lake in 1993 resulted in a homogeneous temperature and oxygen distribution with depth. In spring 1994, the mixing was applied intermittently with a reduction of 75% of the energy costs, while the mixing was still sufficient to prevent stratification. 5. Determination of the buoyancy state of the colonies on a sunny and calm day showed that the buoyancy loss was low close to the bubble plumes, and high at some distance from these plumes. This suggests that Microcystis could escape the mixing at some distance from the plumes, and could synthesize more carbohydrates during its stay in the upper illuminated layer of the lake than the deep mixed colonies close to the bubble plumes. Determination of the buoyancy state appeared to be a good and simple method to investigate the extent of entrainment of colonies in the turbulent flow.

The determination of total nitrogen and total phosphorus concentrations in freshwaters from land use, stock headage and population data: testing of a model for use in conservation and water quality management

Abstract: 1. Nutrient concentrations (particularly N and P) determine the extent to which water bodies are or may become eutrophic. Direct determination of nutrient content on a wide scale is labour intensive but the main sources of N and P are well known. This paper describes and tests an export coefficient model for prediction of total N and total P from: (i) land use, stock headage and human population; (ii) the export rates of N and P from these sources; and (iii) the river discharge. Such a model might be used to forecast the effects of changes in land use in the future and to hindcast past water quality to establish comparative or baseline states for the monitoring of change. 2. The model has been calibrated against observed data for 1988 and validated against sets of observed data for a sequence of earlier years in ten British catchments varying from uplands through rolling, fertile lowlands to the flat topography of East Anglia. 3. The model predicted total N and total P concentrations with high precision (> 95% of the variance in observed data explained). It has been used in two forms: the first on a specific catchment basis; the second for a larger natural region which contains the catchment with the assumption that all catchments within that region will be similar. Both models gave similar results with little loss of precision in the latter case. This implies that it will be possible to describe the overall pattern of nutrient export in the UK with only a fraction of the effort needed to carry out the calculations for each individual water body. 4. Comparison between land use, stock headage, population numbers and nutrient export for the ten catchments in the pre-war year of 1931, and for 1970 and 1988 show that there has been a substantial loss of rough grazing to fertilized temporary and permanent grasslands, an increase in the hectarage devoted to arable, consistent increases in the stocking of cattle and sheep and a marked movement of humans to these rural catchments. 5. All of these trends have increased the flows of nutrients with more than a doubling of both total N and total P loads during the period. On average in these rural catchments, stock wastes have been the greatest contributors to both N and P exports, with cultivation the next most important source of N and people of P. Ratios of N to P were high in 1931 and remain little changed so that, in these catchments, phosphorus continues to be the nutrient most likely to control algal crops in standing waters supplied by the rivers studied.

Inland dispersal of adult aquatic insects

Abstract: 1. Adult caddisflies (Trichoptera) and mayflies (Ephemeroptera) were light-trapped on summer evenings along the Detroit River and Lake St. Clair, near Windsor, Ontario, Canada. Light traps were located at the shore and at increasing distances inland up to 5 km, and were operated simultaneously for 2 h following sunset. Catches of five species of caddisflies of the family Hydropsychidae (Cheumatopsychecampyla, Cheumatopsychespeciosa, Hydropsychehageni, Hydropsychephalerata, Macrostemumzebratum) and the mayfly Hexagenia (Ephemeridae) were used to examine inland distribution. 2. Inland dispersal was limited: catches of caddisflies declined at a greater than exponential rate with increasing distance from shore. Mean dispersal distance from the shoreline ranged from 650 to 1845 m. Smaller caddisfly species dispersed shorter distances than larger caddisflies and Hexagenia. 3. Inland distribution of adult caddisflies exhibited considerable interspecific variation: distribution was inconsistent among trials for Hexagenia, possibly owing to timing of collections in relation to periods of peak emergence. 4. Sex ratios of caddisflies were female biased at most sites. No consistent bias was observed for Hexagenia. Different inland distribution patterns were observed for males and females. The differences appeared to reflect species-specific reproductive strategies.

Oral deformities in tadpoles (Rana catesbeiana) associated with coal ash deposition: effects on grazing ability and growth

Abstract: 1. Tadpoles of the bullfrog (Ranacatesbeiana) collected in a coal ash deposition basin (contaminated with As, Cd, Cr, Cu, Se and other elements) and a downstream drainage swamp had a reduced number of labial teeth and deformations of labial papillae when compared with tadpoles from reference areas. Tadpoles from the coal ash-affected areas had 90% fewer teeth in anterior tooth row number 2 and 40% fewer teeth in posterior row number 1 than reference animals. In the deposition basins, drainage swamp and reference ponds, respectively, 96.2, 85.1 and 2.9% of tadpoles had oral deformities. 2. Tadpoles with deformities were less able to graze periphyton than were normal tadpoles, when tested in the laboratory. When presented with periphyton as a sole food source, tadpoles with deformed teeth had lower (negative) growth rates than those with normal teeth, which had slightly positive growth rates. When particulate food was also available, tadpoles grew well regardless of deformities. 3. It appears that the morphological deformities associated with this coal ash-polluted environment can have ecological ramifications for the affected organisms by limiting the type of food that can be consumed and the ability to grow when multiple food types are unavailable.

Grazing by a native and an exotic crayfish on aquatic macrophytes

Abstract: 1. We compared grazing by native noble crayfish (Astacus astacus) and the exotic signal crayfish (Pacifastacus leniusculus) on seedling or well-established macrophytes. 2. In a pool experiment, seedlings of emergent Scirpuslacustris and floating-leaved Potamogetonnatans were heavily grazed by adult signal crayfish, whereas established plants of the same species sustained only minor damage. 3. In a preference experiment two submerged macrophytes (Charavulgaris and Elodeacanadensis), and both seedlings and established plants of S. lacustris and P. natans, were presented pairwise to signal and noble crayfish. There was no significant difference in preference by the two crayfish species. Chara vulgaris was preferred to all other plants presented, established plants of S. lacustris and P. natans were never preferred, and seedlings of P. natans were preferred to established P. natans. 4. An aquarium experiment was conducted in which the consumption of Chara by signal and noble crayfish was measured in relation to water temperature. Signal crayfish consumed significantly more Chara than noble crayfish, especially at higher temperatures. 5. Our results indicate that the signal crayfish is the more voracious grazer especially at higher temperatures. There may be negative effects on vegetation (emergents and floating-leaved, as well as submerged species) when the signal crayfish is introduced. Chara species are particularly susceptible, since they are preferred by crayfish and the genus includes a large number of rare species. Stocking of crayfish therefore could lead to the decline or removal of submerged species in order of crayfish feeding preferences and could prevent the spread or cause a decline of emergent and floating-leaved vegetation.

Effects of bioturbation by tube‐dwelling chironomid larvae on oxygen uptake and denitrification in eutrophic lake sediments

Abstract: 1. Oxygen uptake and denitrification were determined in two bioturbated sediments from a eutrophic lake in southern Sweden. In laboratory mesocosms, an organic profundal sediment was incubated with Chironomus plumosus L. and a sandy littoral sediment with an organic-rich top layer was incubated with Polypedilum sp. Both species of chironomid are sediment tube-dwelling. 2. Oxygen consumption, expressed per gram of larval dry weight, was enhanced to the same extent by the larvae in both sediments. Measurements of the respiration rate of individual larvae revealed that the respiration per gram dry weight of the smaller Polypedilum sp. was more than three times higher than that of C. plumosus. 3. Denitrification was measured using the ‘nitrogen isotope pairing’ technique. In the organic sediment, denitrification of nitrate from the water phase (dw) and denitrification of nitrate from coupled nitrification (dn) were each correlated with the biomass of C. plumosus. In the sandy sediment, dw was correlated with the biomass of Polypedilum sp., while dn did not show any correlation with Polypedilum sp. 4. Oxygen uptake in the organic sediment was increased by a factor of 2.5, dw 5-fold and dn 2.5-fold at a biomass of 10 g m–2 dry weight of C. plumosus. The same biomass of Polypedilum sp. in the sandy sediment resulted in a 2-fold stimulation of oxygen uptake and a 3-fold stimulation of dw, while dn was not affected. These differences in stimulation between oxygen uptake and denitrification by the larvae in the sediments suggest that the stimulation pattern cannot be explained by simple extension of the sediment surface. The burrows evidently reduce the distance between the nitrate source in the water column and the denitrifiers in the anoxic zones. 5. This study indicates that bioturbation by macrofauna elements can have a great impact on denitrification in lake sediments, and that different organisms can influence nitrogen turnover in specific ways.

Periphyton dynamics in a floodprone prealpine river: evaluation of significant processes by modelling

Abstract: 1 Periphyton chlorophyll a was measured at weekly or 2 weekly intervals from October 1992 to March 1994 at four sites in a Swiss prealpine gravel bed river that was frequently disturbed by unpredictable spates 2 To evaluate the dominant processes that control periphyton biomass, measured data were compared with a set of simulations from an empirical dynamic periphyton model Different combinations of process hypotheses were systematically activated and deactivated in order to assess their importance 3 The simplest model leading to an acceptable agreement with measured data employs a biomass-dependent growth rate, a detachment rate directly proportional to discharge and biomass, and a catastrophic loss rate during bed moving spates Terms describing light or temperature dependence had a minor effect on the model fit 4 The model describes the temporal pattern of the periphyton biomass as a series of growth curves periodically truncated by spates Within the uncertainties of the measurements, mainly caused by the spatial heterogeneity of periphyton, the biomass recovered along deterministic trajectories 5 Sensitivity analyses with respect to model parameters and model structure showed that site-specific model parameters could not be unequivocally determined, and that the model yields similar results with slightly different formulations of processes This indicates that the data base with respect to periphyton biomass was too small for a unique identification of model details but that the main conclusions on the significance of processes did not depend on arbitrary choices of the model formulation

Reconstructing the past density of planktivorous fish and trophic structure from sedimentary zooplankton fossils: a surface sediment calibration data set from shallow lakes

Abstract: 1. As quantitative information on historical changes in fish community structure is difficult to obtain directly from fish remains in lake sediments, transfer function for planktivorous fish abundance has been developed based on zooplankton remains in surface sediment (upper 1 cm). The transfer function was derived using weighted average regression and calibration against contemporary data on planktivorous fish catch per unit effort (PF-CPUE) in multiple mesh size gill nets. Zooplankton remains were chosen because zooplankton community structure in lakes is highly sensitive to changes in fish predation pressure. The calibration data set consisted of thirty lakes differing in PF-CPUE (range 18–369 fish net–1), epilimnion total phosphorus (range 0.025–1.28 mg P l–1) and submerged macrophyte coverage (0–57%). 2. Correlation of log-transformed PF-CPUE, total phosphorus and submerged macrophyte coverage v the percentage abundance in the sediment of the dominant cladocerans and rotifers revealed that the typical pelagic species correlated most highly to PF-CPUE, while the littoral species correlated most highly to submerged macrophyte coverage. Consequently, only pelagic species were taken into consideration when establishing the fish transfer function. 3. Canonical correspondence analysis (CCA) revealed that the pelagic zooplankton assemblage was highly significantly related to PF-CPUE (axis 1), whereas the influence of total phosphorus and submerged macrophyte coverage was insignificant. Predicted PF-CPUE based on weighted average regression without (WA) and with (WA(tol)) downweighting of zooplankton species tolerance correlated significantly with the observed values (r2 = 0.64 and 0.60 and RMSE = 0.54 and 0.56, respectively). A marginally better relationship (r2 = 0.67) was obtained using WA maximum likelihood estimated optima and tolerance. 4. It is now possible, quantitatively, to reconstruct the historical development in planktivorous fish abundance based on zooplankton fossil records. As good relationships exist between contemporary PF-CPUE data and indicators such as the zooplankton/phytoplankton biomass ratio, Secchi depth and the maximum depth distribution of submerged macrophytes, it is now also possible to derive information on past changes in lake water quality and trophic structure. It will probably prove possible further to improve the transfer function by including other invertebrate remains, e.g. chironomids, Chaoborus, snails, etc., and its scope could be widened by including deeper lakes, more oligotrophic lakes, more acidic lakes and lakes with extensive submerged macrophyte coverage (in the latter case to enable use of the information in the fossil record on plant-associated cladocerans).

Interannual variability in phytoplankton biomass and species composition in a subtropical reservoir

Abstract: SUMMARY 1 The interannual variability of the dominant phytoplankton populations is described in a subtropical reservoir in Queensland using weekly data for a 16-year period between 1978 and 1994. North Pine Dam, Brisbane, is in an area characterized by strong interannual variability in rainfall. This variability is linked to El Nino Southern Oscillation (ENSO) events. Between 1978 and 1994 periods of drought (during strong ENSO events) were interspersed by periods of flooding rains. Rainfall on the catchment and temperature and oxygen within the dam showed strong 40-day periodicities which also varied in strength interannually in response to ENSO events. Similar patterns of fluctuations in the 40-day periodicity have been found elsewhere in SE Australia. Seasonal cycles of stratification in the dam were a function of both hydrographic and hydrological events. Intermittent rain storms caused partial turnovers and large outflows. As much as 90% of the dam volume was exchanged in a single flood event. 2 The dominant phytoplankton species were similar to those frequently found in tropical and subtropical lakes and reservoirs. The phytoplankton community switched between cyanobacterial blooms (Cylindrospermopsis, Microcystis) during drought and falling water levels and diatom blooms (Aulacoseira) in response to inflows and seasonal turnovers. There appeared to be a subtle interaction between inflows, water column stability, the periodic overturns and the occurrence of the dominant species. All the dominant species showed long periods (2–4 years) of exponential increase or decrease superimposed on top of the seasonal fluctuations in abundance. These patterns of abundance led to marked interannual variability in the phytoplankton biomass. Climate variability had a major impact on the seasonal and interannual changes of the dominant phytoplankton species. 3 Phytoplankton biomass tended to be depressed for about 3 months after individual storm events but the data also displayed long-term lag effects (2–4 years) which destroyed any significant correlation between water residence time and biomass. Summer maxima of biomass dominated by cyanobacteria disappeared between 1985 and 1990 and were replaced by smaller winter peaks. The data presented here are not capable of unequivocally identifying the precise reason for these longer-term effects. Because of the implications for water quality management in subtropical and tropical reservoirs they warrant further study.

Recolonization by benthic invertebrates after experimental disturbance in a Swiss prealpine river

Abstract: field. 2. Our field experiment compared the structure of the benthic invertebrate community in the prealpine River Necker in north-eastern Switzerland with predictions of the patch dynamics concept about the faunal composition of frequently disturbed streams. We also compared the resistance and resilience of the invertebrates between two sites in the River Necker. A similar substratum composition at both sites, but higher shear stress values both at baseflow and bankfull discharge at site 2, implied a higher disturbance frequency at the latter site. Five patches of stream bed of µ 9m 2 were disturbed by kicking and raking at each site, while five similar areas served as controls. From each plot, six Surber samples were taken: the first immediately after the disturbance, and the following five 1, 3, 6, 10 and 30 days later. 3. Resilience of the total benthic invertebrate fauna was high. The total number of individuals recovered to undisturbed densities within 30 days at site 1 and 6 days at site 2. Taxon richness recovered within 3 days. In accordance with theory, taxa with high recolonization rates made up a major percentage of the total number of individuals, especially in disturbed plots. However, this percentage was lower at site 2 in spite of the higher disturbance frequency at this site. Rhithrogena spp., Leuctra spp. and the Simuliidae recovered faster to undisturbed densities at site 2. In contrast, absolute recolonization rates of these taxa were higher at site 1, where total invertebrate densities were more than twice as high as at site 2. 4. Our results suggest that the time since the last disturbance should be considered as an important factor in studies of benthic invertebrate communities in prealpine rivers, because disturbances can alter the community structure. In frequently disturbed streams, very short sampling intervals may be needed to detect differences in taxon-specific colonization rates. The specific hydrological disturbance regime of such streams is also important, because even within-stream differences in the resilience of the benthic invertebrate community are possible.

Atyid shrimps (Decapoda: Atyidae) influence the spatial heterogeneity of algal communities over different scales in tropical montane streams, Puerto Rico

Abstract: 1. Atyid (Decapoda: Atyidae) shrimps influence the distribution of algal communities over different scales in tropical montane streams of Puerto Rico. Within pools of an atyid-dominated stream, atyid shrimps enhanced patchiness in algal communities along the depth gradient. Algal bands occurred in shallow pool margins where atyids did not forage (< 3 cm below water surface), with significantly greater standing crop, taxon richness, and structural complexity than deeper areas. In deeper water, atyids reduced small-scale patchiness in algal community composition and maintained a low-growing understorey turf dominated by sessile diatoms (Bacillariophyta) and, sometimes, closely cropped, filamentous blue-greens (Cyanophyta). 2. Among pools of the atyid-dominated stream, atyids interacted with light to determine algal patchiness between stream margins and deeper areas. In sunny pools, algal standing crop was 140-fold greater in pool margins than in deeper areas where atyids foraged. In shaded pools, however, standing crop in pool margins was only 5-fold greater than in deeper areas. Effects of light on algal standing crop were greater outside atyid foraging areas than within, indicating that shrimp grazing overrides the positive effects of light. 3. In contrast to the atyid-dominated stream, algal communities in an atyid-poor stream were characterized by a high biomass of loosely attached epipelic diatoms and no depth zonation. Interstream rock and shrimp transplant experiments indicated that atyids significantly reduced algal standing crop and altered community composition on rocks from atyid-poor streams within 24 h. Results support the hypothesis that atyid shrimps play a major role in determining observed interstream differences in algal communities.

Potential fragmentation and loss of thermal habitats for charrs in the Japanese archipelago due to climatic warming

Abstract: 1. The upper thermal limits of the present distributions of two charr species, Dolly Varden, Salvelinus malma, and white-spotted charr, S. leucomaenis, in streams of the Japanese archipelago were examined using groundwater temperature as an index of thermal condition. The lower limits of the altitudinal distributions of Dolly Varden and white-spotted charr were delineated, respectively, by 8 and 16 °C groundwater isotherms. 2. The potential impact of future climatic warming on the geographical distribution, habitat extent and population fragmentation of each species was predicted at both the full archipelago and individual catchment levels. 3. For Dolly Varden, analysis at the full archipelago level indicated a loss of 27.6, 67.2, 79.6 and 89.6% of the current geographical range, respectively, for a 1, 2, 3 and 4 °C increase in mean annual air temperature. The present distribution area of white-spotted charr would likewise reduce by 4.1, 20.5, 33.8 and 45.6%, respectively. 4. Based on the analyses of three individual catchments, one for Dolly Varden and two for white-spotted charr, the lower habitat boundaries for the two charr species could be expected to rise increasingly to higher elevations in each catchment as warming proceeded. As a consequence, there would be large reductions in mean habitat area, with increasing habitat fragmentation followed by localized extinctions of the two species.

The energetic importance of terrestrial arthropod inputs to three warm‐water streams

Abstract: 1. Inputs of terrestrial arthropods (number and mass m–2 d–1) from riparian corridors to three streams representing different orders were highly variable among seasons and sites, with significantly greater (P < 0.05) inputs at the headwater stream during summer months, compared with other sites and seasons. 2. No significant differences in estimates of stream retention of terrestrial arthropods (number and mass m–2 d–1) were observed among sites; however, retention of terrestrial arthropods at all sites was significantly greater during summer months, compared with other periods. 3. The gravimetric proportion of terrestrial arthropods present in the stomachs of redbreast sunfish (Lepomis auritus) and bluegill (L. macrochirus) was equivalent among sites. However, estimates of the dietary importance of terrestrial arthropods at all study sites were significantly greater in the summer, compared with other seasons. 4. Estimates of the potential annual energetic contribution (kJ m–2 d–1) of terrestrial arthropod inputs to the stream system were comparable with published rates of total annual production of aquatic macroinvertebrates in other Virginia headwater streams. 5. Results of this study supported the hypothesis that terrestrial arthropods represented an important energetic subsidy to stream fish during periods of low aquatic macroinvertebrate availability, and suggest that this component of allochthonous input is a potentially significant, but poorly understood energetic linkage between riparian landscapes and stream ecosystems.

Diatoms as indicators of river quality in the Nepalese Middle Hills with consideration of the effects of habitat‐specific sampling

Abstract: 1 Using a replicated survey design at the catchment scale, we compared the composition and diversity of diatom communities in three stream groups in the Middle Hills of Nepal: semi-natural reference (Arun Valley), enriched by agricultural runoff (Likhu Khola) and grossly polluted by sewage (Kathmandu Valley) We sampled riffles at all sites; in the Kathmandu Valley and Likhu Khola we also sampled diatoms in pools and on vegetation 2 Species richness and diversity H′ were significantly higher in agricultural streams than in either organically polluted streams or references Community composition, as shown by principal components analysis, varied significantly between all three stream types due to differences in the abundances of species characteristic of organic enrichment (Kathmandu Valley), moderate enrichment (Likhu Khola) or undisturbed hillstreams (Arun Valley) 3 Few species varied in relative abundance between pools, riffles and vegetation so that variations in community composition and diversity were stronger between stream groups than between habitats Samples from any one habitat produced only 757% (± 86 SD) of the species from three habitats, so that surveys aimed at recording biodiversity may need more comprehensive habitat coverage than surveys for biomonitoring 4 We conclude that diatom communities can indicate different sources of pollution in Nepalese streams, and advocate further studies to develop this indicator potential over a wider area of the Himalaya Comparisons between replicate groups of streams selected a priori helped to clarify effects which were sometimes obscured by other survey designs

The relationships among three habitat scales and stream benthic invertebrate community structure

Abstract: 1. The relationships between three habitat scales and lotic invertebrate species composition were investigated for the 15 540 km2 Yakima River basin in south-central Washington, U.S.A. 2. The three spatial scales were sample (the sampled riffle), reach (a length of ten–twenty stream widths) and segment (a length of stream of nearly uniform slope and valley form having no change in stream order). 3. Physical variables were highly correlated between scales and expressed a relationship between altitude, basin form and small-scale physical structure. 4. Multiple discriminant function analyses indicated that segment- and reach-scale variables discriminated among species-defined groups better than sample-scale variables. 5. Species composition varied along a complex altitudinal gradient of changing basin form and resultant land use. 6. There was no clear relationship between species richness and altitude on a site basis. However, when viewed at the basin scale, maximum richness was observed at the transition between montane and valley sites.

Patch selection by mottled sculpin (Pisces: Cottidae) in a southern Appalachian stream

Abstract: 1. We examined the effects of prey abundance on patch selection by a benthic fish, the mottled sculpin (Cottus bairdi), in a fourth order, southern Appalachian stream (North Carolina, U.S.A.). This habitat is a mosaic of small (i.e. < 0.5 m2), relatively discrete patches. 2. Patches were characterized in terms of physical habitat variables, detritus (coarse particulate organic matter, CPOM) and macroinvertebrate abundance (number and biomass). We quantified patch selection by comparing the characteristics of patches utilized by sculpin with those of locally available patches. Locally available patches were selected using a constrained random sampling design (i.e. randomly selected within a 2 m radius from each fish). We also examined the relationship between macroinvertebrate abundance, CPOM and the physical characteristics of available patches. 3. Patches selected by sculpin contained significantly higher macroinvertebrate abundances (both number and biomass) than locally available patches in five out of six seasonal samples. Sculpin also occupied patches with significantly higher amounts of CPOM in three out of five seasonal samples. Patches utilized by sculpin, however, could not be consistently differentiated from locally available patches on the basis of physical variables. In addition, macroinvertebrate abundance was not consistently related to physical habitat variables or CPOM during the course of the study. 4. Our results suggest that sculpin are able to assess patch quality on the basis of prey abundance and select patches that potentially maximize energy gain. This behaviour may produce an increase in individual fitness, especially when prey distributions are not consistently related to habitat variables. Quantifying patch use in relation to prey abundance may help elucidate the causal factors determining habitat use by benthic fishes in other lotic systems.

Intraguild predation and species exclusions in amphipods: the interaction of behaviour, physiology and environment.

Abstract: 1. Data from field surveys, laboratory experiments and computer simulations of community dynamics revealed that a novel interaction among intraguild predation, physiological adaptation and environment may explain the complex distributions of two putatively competing aquatic amphipods. 2. Gammarus pulex and G. tigrinus both thrive in fresh and oligohaline waters in western Europe. However, the native European G. pulex excludes the invading North American G. tigrinus from freshwaters of relatively low conductivity, whereas the reverse occurs at higher conductivities. Additionally, there is much spatio-temporal fluctuation in the patterns of coexistence of these species. 3. Laboratory experiments in The Netherlands and Ireland revealed that mutual predation of moulting individuals occurred frequently between these species. However, predation frequencies were differentially in favour of G. pulex under the ionic conditions to which this species is physiologically adapted (freshwater). On the other hand, predation was not differential under the ionic conditions to which G. tigrinus is physiologically adapted (oligohaline water). 4. A mathematical model, which extends the logistic equation to include mutual intraguild predation, simulated interactions over a range of values of relevant population parameters. This indicated that G. pulex would be excluded when balanced instantaneous rates of mutual predation were combined with the known greater reproductive output of G. tigrinus. However, this reproductive advantage is overcome by any relatively small bias in the instantaneous rate of predation favouring G. pulex, leading to the exclusion of G. tigrinus. This occurs even when the reproductive advantage to G. tigrinus is relatively large. Moreover, the model generated ‘switches’ in species dominance that are determined by the relative values of reproductive rate and mutual predation. The time taken to ‘switch’ may explain the transient periods of apparent coexistence of these species observed in the field. 5. The complex community dynamics of such species may thus be understood in terms of variation in the intensity of species interactions mediated by behavioural, physiological and environmental factors.

Multi-scale approach to species–habitat relationships: juvenile fish in a large river section

Abstract: 1. Juvenile fish were sampled by electrofishing using the point abundance sampling method from August to November 1991 in the Lower Rhone River. Variations in fish abundance between point samples were partitioned according to three spatial scales of habitat heterogeneity. 2. The compartment scale, differentiating channel and ‘dike fields’ (an area marginal to the main channel partially enclosed by an artificial embankment), was associated with variation in water current, temperature and biological productivity. Every juvenile fish species studied was more abundant in the dike fields, using them as a nursery. 3. Within both compartments (i.e. main channel and dike fields), the same five bank types were represented, defining the intermediate scale of habitat heterogeneity. In the dike fields, beaches were selected by most of the species, whereas steep silt banks were more often selected in the main channel. 4. After having removed the variation explained at the two larger habitat scales, only depth and shelter were found to have a significant effect on fish abundance at the microhabitat scale. 5. When compared with a classical single-scale approach, the multi-scale approach was found to explain a greater amount of fish abundance variations and to facilitate the interpretation of observed patterns by scaling the species’ responses to habitat heterogeneity. The difference between the amounts of variation explained was proposed as a measure of how much habitat selection is hierarchical.

Spatial and temporal variation of microbial respiration rates in a blackwater stream

Abstract: 1. The extent of spatial and temporal variation of microbial respiration was determined in a first-order, sand-bottomed, blackwater stream on the coastal plain of south-eastern Virginia, U.S.A. 2. Annual mean respiration rates (as g O2 m–3 h–1) differed significantly among substrata: leaf litter, 12.9; woody debris, 2.4; surface sediment, 0.8; hyporheic sediment, 0.4; water column, 0.003. Rates associated with wood were higher than those with leaves when expressed per unit surface area. 3. Highest respiration rates on leaves, wood and in the water column occurred during the summer, whereas rates in the sediments were greatest during the late autumn and winter. Water temperature, as well as particulate organic matter and nitrogen content of the substrata, was correlated positively with respiration rates. 4. A stepwise multiple regression showed that temperature and nitrogen content together explained 88% of the variation in respiration rates of leaves and wood. In contrast, particulate organic matter content and nitrogen content explained 89–90% of the variation in respiration in the sediments. Although water temperature was a significant factor in the sediment multiple regressions, its addition as an independent variable improved the regression models only slightly. 5. Annual mean respiration in the stream channel, based on the proportional amount of respiration occurring associated with each type of substratum during each month, was 1.1 kg O2 m–2 yr–1. Seventy per cent of respiration in the stream occurred in the hyporheic zone, 8–13% occurred in the surface sediment, leaf litter or woody debris, and < 1% occurred in the water column. Approximately 16% of total detritus, or 40% of non-woody detritus, stored in the stream during the year was lost to microbial respiration.

Amphibian species richness and distribution in relation to pond water chemistry in south‐western Ontario, Canada

Abstract: 1. We assessed the patterns of amphibian species richness and distribution in relation to water chemistry over a large geographical area in 1992–94. 2. Thirteen amphibian species were observed at 180 ponds, with mean species richness 3.5 ± 0.13 species per pond (range zero to nine). Water samples were collected from 143 ponds, analysed for fifteen chemical variables, and further analysed by multivariate statistical techniques. 3. Water in the study area was hard, alkaline and well-buffered against pH change, and most ponds were eutrophic. Amphibian species richness was negatively correlated with five chemical variables (chloride, conductivity, magnesium, total hardness, turbidity). 4. Principle components analysis reduced the data set to four chemical components that explained 65.4% of the variance in the original variables. Principle component scores were retained for use in further multivariate tests. Multiple regression accounted for only 19.0% of the variance in amphibian species richness. Discriminant function analysis (DFA) was used to determine if water chemistry variables discriminated among species, but it was only able to classify 17.5% of cases correctly. DFA was also used to determine if water chemistry distinguished between used and unused sites for individual species. DFA was moderately successful, classifying 61–77% of cases correctly. 5. General water chemistry appears to play only a minor part in affecting amphibian species richness in south-western, Ontario. However, chemical variables may be helpful to distinguish between used and unused sites for some species.

Litter breakdown in streams of the Agüera catchment: influence of dissolved nutrients and land use

Abstract: 1. The breakdown of oak (Quercus robur L.), chestnut (Castanea sativa Miller) and eucalypt (Eucalyptus globulus Labill.) litter enclosed in 5-mm mesh bags was compared between first-order headwaters (two with native riparian forest and two with eucalypt plantations) and a third-order reach of Aguera stream. Weight loss and dynamics of phosphorus and nitrogen in litter were studied for a period of 155 days. 2. Among the different sites, processing rates ranged from 0.0045 to 0.0080 day–1 for chestnut leaf litter, from 0.0036 to 0.0051 day–1 for oak, and from 0.0027 to 0.0158 day–1 for eucalypt. 3. The availability of nutrients in water clearly influenced nitrogen and phosphorus dynamics in litter. In headwater reaches, net immobilization was not observed and losses of phosphorus and nitrogen followed mass loss. However, there was an enrichment of litter at the low reach, where influence of human settlements—located upstream—could lead to a greater availability of phosphorus in water. 4. The enhancement of litter decay by the exogenous nutrient supply depended on leaf quality, as only the processing rate of eucalypt increased at the nutrient-rich site. 5. The processing rates differed little among headwaters, suggesting that riparian forest type, i.e. deciduous forest v eucalypt plantations, did not affect litter decay in the stream.

Differential behavioural responses of mayflies from streams with and without fish to trout odour

Abstract: 1. In streams, mayflies (Order Ephemeroptera) are at risk from fish feeding visually in the water column. The effect of fish odour on the behaviour of Baetis bicaudatus from a fishless stream and a trout stream was investigated in four large oval tanks supplied with water from the fishless stream. 2. For each mayfly population, mayfly positioning on the substratum and movement in the water column (drift) were measured during the day and night, over 3 days. Brook trout (Salvelinus fontinalis) odour was added to two tanks to test the effect of a threat from fish. 3. Throughout the experiment more mayflies from the trout stream were observed on the substratum surface and in the water column during the night than the day, but the magnitude of night drift was less in tanks with fish odour. 4. Baetis from the fishless stream also displayed a nocturnal periodicity in drift and positioning, but their night-time drift was not affected by the presence of fish odour. On the first day of the experiment, however, more mayflies were observed on the substratum surface and drifting in tanks without fish odour during the day. 5. Sensitivity to fish odour may enable mayflies to alter their behaviour according to the risk of predation from fish.

Genetic differentiation among populations of Caridinazebra (Decapoda: Atyidae) in tropical rainforest streams, northern Australia

Abstract: 1. Caridina zebra is a common atyid shrimp in some tropical rainforest streams in far north Queensland, Australia. Genetic variation at five allozyme loci was used to estimate the level of dispersal among populations of this species, within and between stream systems. Shrimps were sampled from nine streams in the Tully River catchment and two headwater streams in the adjacent Herbert River catchment in an area under consideration for extensive hydroelectric development. 2. High levels of genetic differentiation were recorded among most populations which suggests that, like other fully aquatic species, movement is limited to a very small spatial scale. 3. In the Tully catchment, populations of shrimp from streams with confluences at high altitude showed less genetic differentiation than those from streams which directly entered the lower river. Dispersal between the latter streams is clearly limited by the presence of large waterfalls and cascades. 4. Adjacent stream populations were often highly differentiated, despite their close proximity, suggesting that overland dispersal is unlikely. However, populations of shrimp in the two streams in the Herbert catchment were strikingly similar in genetic structure to those in adjacent headwater streams of the Tully. Such similarity may reflect relatively recent changes in drainage patterns.

Mesocosm experiments on the interaction of sediment influence, fish predation and aquatic plants with the structure of phytoplankton and zooplankton communities

Abstract: 1. Little Mere, U.K., received large quantities of sewage effluent until 1991, when the effluent was diverted. Experiments, carried out in mesocosms in 1992 and 1993, were designed to predict the effects of: (i) reduced external nutrient loading; (ii) reduced internal loading from the sediment; and (iii) recolonization by fish of the better aerated water. Treatments included isolation of the water from the underlying sediment or exposure to the sediment (which lacked plants in 1992, but was covered by Potamogeton berchtoldii in 1993) crossed with different population densities of Rutilus rutilus in 1992 and of Perca fluviatilis in 1993. 2. Exposure to sediment (as opposed to isolation from it) resulted in no net change in the biovolumes of most major algal groups, but this masked major complementary effects on individual species. The experiments showed a decreasing influence of the sediment, between 1992 and 1993, in determining water chemistry, and an increasing pH between years but no increase in cyanophyte dominance. This had been anticipated because a lake upstream provides abundant inocula, and conditions in Little Mere after diversion of effluent were expected to favour cyanophytes. 3. Roach and perch additions to the mesocosms resulted in reductions in Daphnia populations but increases in numbers of small Cladocera and copepods. Plant-associated Cladocera were unaffected by fish. The presence of submerged plants to some extent reduced fish predation effects on Daphnia hyalina. 4. The experimental results in general accurately predicted subsequent events in the open lake.