The effect of the release of an artificial discharge of water on invertebrate drift in the R. Wye,Wales

An artificial discharge of water (3.0 m³/sec), over a 48 h period, from an impoundment into the R. Wye did not substantially affect water temperature or concentrations of dissolved oxygen and suspended solids at a site 16 km below the impoundment. However, the load of suspended material on the second day of the release was about 10 times greater than the pre-release load. The total number of drifting macroinvertebrates on the first and second days of the release were about 7 and 3 times greater than the number on the day preceding the release. The initial increase in flow at 15.00 h resulted in an immediate increase in the number of drifting larvae of Rheotanytarsus, a tubicolous chironomid. Subsequently there was an enhanced night-time increase in the total number of drifting invertebrates, particularly the mayfly, Ephemerella ignita (Poda), and this also occurred on the second night of the release. Increases in the number of drifting Rheotanytarsus and Ephemerella, the most abundant invertebrates, resulted in increase in drift density.Authors' address Llysdinam Field Centre, UWIST, Newbridge-on-Wye, Llandrindod Wells, Powys, Wales.     

A comparison of the relative contributions of temporal and spatial variation in the density of drifting invertebrates in a Dorset (U.K.) chalk stream

1. Invertebrate drift is commonly investigated in streams, with the majority of studies focussed on temporal (typically diel) variation. In comparison, few studies have investigated spatial variation in drift and there is little consensus among them. We tested the hypothesis that spatial variation in invertebrate drift is as important as temporal variation. 2. The density of drifting invertebrates in a chalk stream was sampled using an array of nets arranged to determine vertical, lateral and longitudinal variation. Samples were collected at dawn, during the day, at dusk and by night, on four separate monthly occasions. Insecta and Crustacea were analysed separately to identify the effect of differing life history strategies. The density of drifting debris was also recorded, to act as a null model. 3. Time of day and vertical position together explained the majority of the variance in invertebrate drift (79% for Insecta and 97% for Crustacea), with drift densities higher at dusk and night, and nearer the stream bed. Independently, time of day (38%, Insecta; 52%, Crustacea) and vertical position (41%, Insecta; 45%, Crustacea) explained a similar amount of the observed variance. Month explained some of the variance in insect drift (9%) but none for Crustacea. 4. Variation in the density of drifting debris showed little in common with invertebrate drift. There was little variation associated with time of day and only 27% of the observed variation in debris could be explained by the factors investigated here, with month explaining the largest proportion (20%). We suggest the difference in drifting debris and invertebrates provides further evidence for a strong behavioural component in invertebrate drift. 5. Spatial variation in invertebrate drift can be of the same order of magnitude as the much‐described diel temporal variation. The extent of this spatial variation poses problems when attempting to quantify invertebrate drift and we recommend that spatial replication should be incorporated into drift studies.     

Diel,size-differential drift patterns of three macroinvertebrate species in the lower Mississippi River,Louisiana (USA)

We explored macroinvertebrate size-differential drift in the lower Mississippi River (a 9th order system). Because this river system is highly turbid, we hypothesized that visually-dependent vertebrate predators feeding on drifting organisms would be at a disadvantage. Thus, size-differential drift should not occur. For one 24-hour period in both January and April, six drift nets were used to sample surface drift. Nets were emptied once every four hours. Individual intra-ocular distances of three macroinvertebrate species (Hydropsyche orris: Trichoptera, Hexagenia limbata: Ephemeroptera, Macrobrachium ohione: Crustacea) were measured. Percentages of size classes in the drift were determined. In both months, large individuals of H. orris and H. limbata were prevalent in the nocturnal but scarce in the diurnal drift. In January, large M. ohione drifted regardless of time. In April, large M. ohione predominated the nocturnal drift. Our results could not be attributed solely to vertebrate predator avoidance. Other mechanisms such as diel microhabitat migration and current velocity may have accounted for the results.     

Diel drift of Chironomidae larvae in a pristine Idaho mountain stream

Simultaneous hourly net collections in a meadow and canyon reach of a mountain stream determined diel and spatial abundances of drifting Chironomidae larvae. Sixty-one taxa were identified to the lowest practical level, 52 in the meadow and 41 in the canyon. Orthocladiinae was the most abundant subfamily with 32 taxa and a 24 h mean density of 294 individuals 100 m⁻³ (meadow) and 26 taxa and a mean of 648 individuals 100 m⁻³ (canyon). Chironominae was the second most abundant subfamily. Nonchironomid invertebrates at both sites and total Chironomidae larvae (meadow) were predominantly night-drifting. Parakiefferiella and Psectrocladius were day-drifting (meadow) whereas 8 other chironomid taxa (meadow) and 2 taxa (canyon) were night-drifting. All others were aperiodic or too rare to test periodicity, Stempellinella cf brevis Edwards exhibited catastrophic drift in the canyon only. The different drift patterns between sites is attributed to greater loss of streambed habitat in the canyon compared to the meadow as streamflow decreased. Consequent crowding of chironomid larvae in the canyon caused catastrophic drift or interfered with drift periodicty. This study adds to knowledge of Chironomidae drift and shows influences on drift of hydrologic and geomorphic conditions.     

Drift of stream invertebrates below a cave source

Stream invertebrate drift below the cave source of South Branch Creek, Minnesota, generally increased rapidly. Ephemeroptera and Trichoptera increased rapidly and then decreased at the lower stations (to 430 m) below the cave. Chironomidae drift, in high numbers but low biomass because of their small size, increased fairly rapidly and leveled out at the lowermost stations. Gastropods increased slowly below the cave, reached a maximum, then decreased somewhat at the lowermost station. Drifting oligochaetes, small in size but in very large numbers, increased more slowly below the cave and appeared not to have reached a maximum at the lowermost station. It was concluded that, in general, drift increases fairly rapidly below the stream origin and fluctuates in the upper reaches, probably reflecting benthic population abundance and local ecological conditions, before attaining equilibria downstream, rather than increasing linearly due to cumulative effects. Paper no. 11,172 Scientific Journal Series, Minnesota Agricultural Experiment Station, St. Paul, Minnesota 55108 Paper no. 11,172 Scientific Journal Series, Minnesota Agricultural Experiment Station, St. Paul, Minnesota 55108     

The effect of long pools on the drift of macro-invertebrates in a mountain stream

Macro-invertebrate drift was measured entering and leaving two pools on the Middle Fork of the Cosumnes River, a third order California stream. Drift rates for Baetis spp., Chironomidae, Simulium spp., Capniidae and total drift were calculated. Significant differences in the numbers of organisms entering the two pools were found for Baetis, Chironomidae, and Capniidae. Comparisons of drift rates at the upstream and downstream ends of each pool showed that the abundance of Chironomidae, Simulium, Capniidae and total drift changed in different directions across the pools. The numbers of organisms leaving the two pools, however, were not significantly different for Baetis, Simulium, Capniidae and total drift. These findings lead us to hypothesize that long pools act as barriers, not filters, to stream macro-invertebrate drift. The composition of drift leaving the pools in this experiment appeared to be controlled by the composition of the benthic habitat at the tail of the pool and not by the composition of upstream drift entering the pools.     

Influence of a metal-contaminated tributary on the invertebrate drift fauna of the King River (Tasmania,Australia)

Drifting invertebrates were collected upstream of, at, and downstream of the input of metal contaminated water into the previously unpolluted King River in western Tasmania. In the upstream section the drift fauna showed characteristics similar to those observed by other workers with a marked nocturnal peak. Changes in drift rates at the site of input were observed and it is hypothesized that mayfly nymphs, and possibly other animals, respond to the change in water quality by leaving the water column and randomly searching the substrate for clean water. Most nymphs re-enter the drift in a current of clean water which was displaced towards the opposing bank by the entry of contaminated water. This re-entry of animals into the drift occurs some four hours after the main nocturnal peak in drift rates.     

A comparative study of the dispersal of 10 species of stream invertebrates

1. Apart from downstream dispersal through invertebrate drift, few quantitative data are available to model the dispersal of stream invertebrates, i.e. the outward spreading of animals from their point of origin or release. The present study provides comparative data for 10 species, using two independent methods: unmarked animals in six stream channels built over a stony stream and marked animals in the natural stream. Experiments were performed in April and June 1973 and 1974, with initial numbers of each species varying from 20 to 80 in the stream channels and 20 to 60 for marked animals. 2. Results were the same for marked invertebrates and those in the channels. Dispersal was not density‐dependent; the number of dispersing animals was a constant proportion of the initial number for each species. The relationship between upstream or downstream dispersal distance and the number of animals travelling that distance was well described by an inverse power function for all species (exponential and log models were poorer fits). Results varied between species but were similar within species for the 4 months, and therefore were unaffected by variations in mean water velocity (range 0.04–0.35 m s^?1) or water temperature (range 6.7–8.9 °C in April, 12.1–14.8 °C in June). 3. Species were arranged in order, according to their dispersal abilities. Three carnivores (Perlodes, Rhyacophila, Isoperla) dispersed most rapidly (70–91% in 24 h, maximum distances 9.5–13.5 m per day), followed by two species (Protonemura, Rhithrogena) in which about half their initial numbers dispersed (50–51% in 24 h, 7.5–8 m per day), and four species (Ecdyonurus, Hydropsyche, Gammarus, Baetis) in which less than half dispersed (33–40% in 24 h, 5.5–7 m per day). Dispersal was predominantly upstream for all nine species. Few larvae (20%) of Potamophylax dispersed, with similar maximum upstream and downstream distances of 3.5 m per day. The mean time spent drifting downstream was known for seven species from previous studies, and correlated positively with their dispersal distances. Therefore, the species formed a continuum from rapid to very slow dispersers. These interspecific differences should be considered when evaluating the role of dispersal in the maintenance of genetic diversity in stream invertebrates, and in their ability to colonise or re‐colonise habitats.     

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).     

Upstream-downstream movements of aquatic invertebrates in a Rocky Mountain stream

Simultaneous collections of drift and organisms moving either upstream or downstream in association with the substrate were made using a specially designed sampler. Samples were taken in a diel series along a transect across the study riffle of a Colorado foothills stream on six dates over an annual cycle. In addition to longitudinal movements, taxonomic composition and diel periodicity were evaluated. The insect-dominated fauna showed a net downstream displacement. Only the caddisflies Helicopsyche borealis and Hesperophylax occidentalis exhibited net upstream movement, primarily a result of low drift frequencies. The taxonomic composition of moving invertebrates differed from that of the benthos. Drift resembled downstream moving substrate-associated invertebrates in composition, but differed from that of the upstream directed fauna. Taxa collectively exhibited four types of diel patterns: 1) similar downstream (drift and substrate-associated movements) patterns, which generally differed from the upstream pattern; 2) similar benthic (upstream and downstream) patterns, which differed from that of drift; 3) aperiodic patterns; and 4) independent patterns for each type of directional movement. Analysis of size classes based on head capsule width for the mayfly Baetis tricaudatus showed significantly smaller size in stationary individuals compared with moving individuals in the population and revealed that nymphs moving during the day were smaller than those moving at night.     

Time spent in the drift by downstream-dispersing invertebrates in a Lake District stream

1. The objective was to determine the time spent in the drift by different taxa of stream invertebrates. Most data were obtained from an earlier experimental study to determine the distances travelled by drifting invertebrates of 16 taxa in Wilfin Beck. Experiments were performed at two sites: ‘site 4’ in a stony, fast‐flowing, section of stream, ‘site 3’ in a deeper stream section where macrophytes were abundant. 2. The significant relationship between the mean distance x (m) travelled in the drift and modal water velocity V (m s^?1) was described by a power function in the earlier study but, as the power was close to one, a linear relationship has now been found to provide a satisfactory model. The rate of increase in x (m) with increasing V varied considerably between taxa. The mean time [mean t (s)] spent in the drift was estimated by dividing each x (m) by the appropriate V. Mean t (s) for each taxon was usually very constant over a wide range of V at each site (0.10–0.60 m s^?1 at site 4, 0.15–0.53 m s^?1 at site 3). A simple model estimated the time spent in the drift by different percentages (e.g. 75, 50, 10 and 1%) of the drifting invertebrates. 3. The experimental taxa at site 4 were divided into three groups according to the mean time spent in the drift. Mean t (s) for the five taxa in group 1 (32.8 s) was not significantly different from that obtained in control experiments with a mixed group of dead invertebrates. A similar time (33.0 s) was obtained for the five taxa in group 2, except at water velocities less than 0.2 m s^?1 when the mean t (s) decreased to 15–21 s. Mean t (s) was constant for each of the six taxa in group 3, and significantly less than that for groups 1 and 2. Mean values ranged from 28.8 s for Ephemerella ignita to only 9.4 s for Baetis rhodani and Gammarus pulex. All mean values were lower at site 3, presumably because of the dense stands of macrophytes, with mean values of 12.9 s for the five taxa in group 1 (equalling the value for dead invertebrates). Mean values for the six remaining taxa varied from 6.4 s for Simulium spp. to only 4.9 s for Baetis rhodani and 4.8 s for Gammarus pulex. It was concluded from a discussion of this study that the time spent in the drift may provide a useful measure for comparing the downstream dispersal of invertebrates in different streams, and may be a useful addition to models for the drift feeding of salmonids.     

The Epipelic Algal Flora of the River Wye System,Wales, U. K. 2. Algal Phyla and Species Population Dynamics

The Bacillariophyta dominated over the other phyla and were mainly recorded in high densities during summer and autumn. The Chlorophyta and Myxophyta (Cyanobacteria) were represented by coccoid forms and by non-heterocystous, filamentous forms, respectively. Both were mainly recorded during summer and mostly absent during winter. Other phyla were occasionally recorded in low densities; the Chrysophyta being found in the River Elan and at one site on the River Wye during June 1980. Their members were considered as “contaminants” or “fall-out” from other communities. Pennate diatoms were the most “constant” species and either showed a general upstream or downstream increase or a general distribution throughout the study area. Populations of the same species colonising both sediments and stones were not correlated or insignificantly correlated with each other at most stations. It was concluded that sediments were unsuitable for algal colonisation. The River Elan and upper Wye were rated as oligosaprobic, the River Ithon and lower Wye as β-mesosaprobic and the River Lugg as α-mesosaprobic.     

The costs of becoming nocturnal: feeding efficiency in relation to light intensity in juvenile Atlantic Salmon

1. Most animals are active by day or by night, but not both; juvenile salmonids are unusual in that they switch from being predominantly diurnal for most of the year to being nocturnal in winter. They are visual foragers, and adaptations for high visual acuity at daytime light intensities are generally incompatible with sensitive night vision. Here we test whether juvenile Atlantic Salmon Salmo salar are able to maintain their efficiency of prey capture when switching between diurnal and nocturnal foraging.
2. By testing the ability of the fish to acquire drifting food items under a range of manipulated light intensities, we show that the foraging efficiency of juvenile salmon is high at light intensities down to those equivalent to dawn or dusk, but drops markedly at lower levels of illumination: even under the best night condition (full moon and clear sky), the feeding efficiency is only 35% of their diurnal efficiency, and fish will usually be feeding at less than 10% (whenever the moon is not full, skies are overcast or when in the shade of bankside trees). Fish were unable to feed on drifting prey when in complete darkness.
3. The ability of juvenile salmon to detect prey under different light intensities is similar to that of other planktivorous or drift-feeding species of fish; they thus appear to have no special adaptations for nocturnal foraging.
4. While winter drift abundance is slightly higher by night than by day, the difference is not enough to compensate for the loss in foraging efficiency. We suggest that juvenile salmon can nonetheless switch to nocturnal foraging in winter because their food requirements are low, many individuals adopting a strategy in which intake is suppressed to the minimum that ensures survival.     

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.     

Drift patterns in a high Andean stream

The drifting of invertebrates was sampled for six 24-hour periods from September 1976 to July 1977 in a small stream of the paramo of the Ecuadorian Andes. The composition of the drift is similar to that of the benthos, though percentages may differ markedly. Drift is relatively constant throughout the year, except in March when unusually high rates were noted; at this same period a marked reduction in the benthos was also observed. Diel periodicities in the drift are unclear, although on the whole drift is more important during the daytime; when individual data series are analyzed, weak patterns of day or night drifting can be recognized in some groups. The input of insects through drift from the small streams is thought to be an important source of food for the salmonid fish inhabiting the larger torrents.     

The drift distances and time spent in the drift by freshwater shrimps, Gammarus pulex, in a small stony stream, and their implications for the interpretation of downstream dispersal

1. The objectives were (i) to determine experimentally and to model the relationship between mean water velocity and both the mean distance travelled, and the mean time spent, in the drift by freshwater shrimps, Gammarus pulex; (ii) to develop a drift distance–water velocity model from the experimental study, and validate it with field data; (iii) to examine the relationship between drift rate, water velocity and benthic density with the latter expressed as a mean value for the whole stream and a mean value corrected for the distance travelled in the drift. 2. In field experiments at 10 water velocities (0.032–0.962 m s^?1), the significant relationship between the mean drift distance and mean water velocity was described both by a power function (power, 0.96) and a linear relationship. The mean drift time was fairly constant at 8.3 s (95% CL ± 0.4). A simple model estimated the drift distance and time spent in the drift by different percentages of the drifting invertebrates. This model predicted correctly the positive relationship between drift rate and water velocity for field data over a year. 3. The relationship between drift rate per hour and the independent variables, water velocity and benthic density, was well described by a multiple‐regression model. Adding temperature and date did not improve model fit. Variations in water velocity and benthic density explained 96% of the variation in nocturnal drift rate (65% to velocity, 31% to benthic density), but only 40% of the variation in diurnal drift rate (29% to velocity, 11% to benthic density). Correcting benthic density for the drift distances did not improve model fit. 4. The significance of this study is that it developed models to predict drift distances and time, values being similar to those obtained in another, larger stream. It also illustrated the importance of spatial scale in the interpretation of drift by showing that when drift distances were taken into account, the impact of drift on the population was higher (4–10% lost day^?1) than when drift distances were ignored (usually < 3% lost day^?1), especially at a local level.