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.     

An epidemic model in a patchy environment

An epidemic model is proposed to describe the dynamics of disease spread among patches due to population dispersal. We establish a threshold above which the disease is uniformly persistent and below which disease-free equilibrium is locally attractive, and globally attractive when both susceptible and infective individuals in each patch have the same dispersal rate. Two examples are given to illustrate that the population dispersal plays an important role for the disease spread. The first one shows that the population dispersal can intensify the disease spread if the reproduction number for one patch is large, and can reduce the disease spread if the reproduction numbers for all patches are suitable and the population dispersal rate is strong. The second example indicates that a population dispersal results in the spread of the disease in all patches, even though the disease can not spread in each isolated patch.     

Transverse mixing of transported material in simple and complex stream reaches

Stream ecologists often make assumptions about the extent to which transported materials (such as dissolved nutrients, particulate organic matter and pollutants) are mixed across the width of a channel. Transverse spatial heterogeneity arises whenever material enters the channel at a point or restricted source, and is dissipated downstream as mixing evens out concentration gradients. A frequent assumption is that the distance required for complete mixing is negligibly short, and therefore that transverse heterogeneity in transported material is not to be expected. However, few mixing data are available for small (discharge <5 m³/s) channels. We measured mixing distances in three small (0.04–0.1 m³/s) eastern Iowa streams via mid-channel injections of a fluorescent tracer. Mixing distances varied, with effectively complete mixing (to statistically undetectable heterogeneity in tracer concentration) taking as little as 5 m or as much as 100⁺ m. Mixing was more rapid in more complex stream reaches and across constrictions and pools (vs. simpler channels). These mixing distances are not negligible when compared to typical spatial scales for ecological experiments.     

Inverse density dependent parasitism in a patchy environment: a laboratory system

Abstract. 1. Two species of parasitoids (Anisopteromalus calandrae (Howard) and Heterospilus prosopidis Vier) attacking the bruchid beetle, Callosobruchus chinensis (L.), show marked inverse density dependent relationships between per cent parasitism and host density per patch.
2. These patterns are well described quantitatively using data on the spatial distribution of searching time by the parasitoids and their attack rates on patches of different host density.
3. A model of optimal foraging predicts just the opposite (i.e. density dependent) patterns of parasitism.
4. Both density dependent and inversely density dependent spatial patterns of parasitism can be explained mechanistically in terms of (a) the allocation of searching time in patches of different host density and (b) the maximum attack rate per parasitoid that constrains the extent of host exploitation within a patch.     

The Ross-Macdonald model in a patchy environment

We generalize to n patches the Ross-Macdonald model which describes the dynamics of malaria. We incorporate in our model the fact that some patches can be vector free. We assume that the hosts can migrate between patches, but not the vectors. The susceptible and infectious individuals have the same dispersal rate. We compute the basic reproduction ratio R(0). We prove that if R(0)1, then the disease-free equilibrium is globally asymptotically stable. When R(0)>1, we prove that there exists a unique endemic equilibrium, which is globally asymptotically stable on the biological domain minus the disease-free equilibrium.     

Drift and settlement of stream insects in a complex hydraulic environment

1. The hydraulic and geomorphic characteristics of stream patches are often associated with distinctive assemblages or densities of stream invertebrates, and it is routinely presumed that these patterns reflect primarily species‐specific habitat requirements. An alternative hypothesis is that such patterns may be influenced by constraints on movement, such as the results of departure and settlement processes. We describe a manipulative experiment that examined how the hydraulic environments created by topographic bedforms influenced the drift behaviour and potential settlement sites for two species of mayfly (Baetis rhodani and Ecdyonurus torrentis). These species are common in the drift and often co‐occur in streams, but differ in their small‐scale distribution patterns, body shape and movement behaviour. 2. Flume experiments were carried out to determine how the hydraulic environments conditioned by a step bedform influence the behaviour of mayflies in the drift (swimming, posturing, tumbling), and the consequences of those behaviours (drift distance and time), compared to drift over a plane bed. The ramped step in the flume mimicked step bedforms that are common in coarse‐grained, high‐gradient streams. In contrast to the plane bed, a zone of recirculating flow was created downstream of the step, above which flow was faster and more turbulent. Uniform flows are used in most flume studies of drift; our approach is novel in recreating a complex hydraulic environment characteristic of stream channels. 3. Both species had some behavioural control over drift, and drift distances and times were shorter for live larvae than for dead larvae over the plane bed. The step had no impact on drift time or distance for live Baetis, but dead larvae were trapped in the flow separation eddy and drift time increased accordingly. Some Ecdyonurus also became trapped in the eddy, but live larvae drifted farther than dead larvae, and farther over the step than the plane bed. 4. Whilst in the drift, larvae altered their behaviour according to the ambient hydraulic environment, but in a species‐specific manner. Over the plane bed, Baetis had occasional swimming bursts, but primarily postured (maintained a stable body orientation), whereas Ecdyonurus spent roughly equal time posturing and swimming. In the more turbulent flows generated by the step, Baetis spent proportionately more time swimming, whereas Ecdyonurus spent more time posturing and often tumbling as body orientation became unstable. 5. In a high‐gradient stream, Baetis was more abundant close to steps than in plane bed patches with less complex flow, whereas the opposite pattern held for Ecdyonurus. Thus, the small‐scale distribution patterns of these species within streams correspond to their drift behaviours and ability to access various hydraulic patch types in our flume. These results are consistent with the hypothesis that constraints on movement and settlement may be important driver of distribution patterns within streams.     

Factors influencing nitrate depletion in a rural stream

A mass balance procedure was used to analyze rates of nitrate depletion in three adjacent reaches of West Duffin Creek, Ontario, Canada. Daily nitrate losses in individual reaches were highly variable (0.5–24 kg N) during low and moderate stream flows in May–October, 1982–1985. Nitrate removal efficiency (nitrate loss as a % of nitrate input) showed a rapid exponential decline with increased nitrate inputs to each reach. Nitrate losses and nitrate removal efficiency also had a significant negative correlation with stream discharge. The association of large nitrate loads with high stream discharge reduced the nitrate removal capacity of the stream because of shorter residence times and a higher ratio of water volume to stream bed area. Water temperature exhibited a significant positive correlation with nitrate loss which may reflect increased denitrification at higher temperatures.Variations in nitrate losses and nitrate removal efficiency between the three reaches were highly influenced by differences in water residence time. Standarized nitrate losses with respect to water residence time revealed a longitudinal decline in nitrate depletion between the reaches which was associated with a downstream decrease in stream nitrate concentration and in the organic carbon content of fine textured sediments from pool habitats.     

Invertebrate drift and benthic community dynamics in a lowland neotropical stream, Costa Rica

In this study we quantified invertebrate drift and related it to the structure of the benthic community, over a 6–8 month period, in a 4th-order tropical stream in Costa Rica. Relative to reports from similar-sized temperate and tropical streams, drift densities were high (2-fold greater: mean 11.2 m⁻³; range 2.5–25 m⁻³), and benthic insect densities were relatively low (>3-fold lower: mean 890 m⁻²; range 228–1504 m⁻²). Drift was dominated by larval shrimps that represented more than 70% of total drift on any given date; the remaining 30% was composed of 54 insect taxa. Among insects, Simuliidae and Chironomidae (Diptera) and Baetidae, Leptohyphes and Tricorythodes (Ephemeroptera) comprised 24% of total drift. Drift periodicity was strongly nocturnal, with peaks at 18:00 h (sunset) and 03:00 h. Our results, and those of previous experiments in the study stream, suggest that nighttime drift is driven by the presence of predatory diurnal drift-feeding fishes and nocturnal adult shrimps. There were no clear seasonal patterns over both ‘dry’ and wet seasons, suggesting that benthic communities are subject to similar stresses throughout the year, and that populations grow and reproduce continuously. This revised version was published online in July 2006 with corrections to the Cover Date.     

Dispersal and recruitment of fish in an intermittent stream network

Animal movement is an important process connecting habitats in heterogeneous landscapes, and can play a key role in population persistence. Laboratory swim trials were conducted to determine and compare the dispersal capabilities of two native Australian fish, mountain galaxias (Galaxias olidus, Family Galaxiidae) and southern pygmy perch (Nannoperca australis, Family Nannopercidae) that maintain populations in hydrologically variable and intermittently flowing streams in south‐eastern Australia. These experiments showed that G. olidus had significantly greater swimming endurance under a range of flow velocities. Concurrent field surveys were used to establish whether swimming abilities observed in laboratory studies were consistent with patterns of inferred movement from distribution and abundance patterns observed in the field. Data collected at multiple sites from headwater to lowland reaches along multiple streams revealed substantial temporal changes in the distribution of young‐of‐year (0+) G. olidus, with spawning occurring at upland sites in winter, followed by downstream larval migration and subsequent upstream movement in late spring. Observed spatial and temporal patterns in G. olidus abundances were consistent with a source‐sink population structure, which may be disrupted by prolonged cease‐to‐flow periods during drought years. In contrast, results for N. australis suggested limited dispersal, with restricted local populations that persist at sites with permanent surface water. These field and laboratory findings complement our understanding of the spatial population structure of these two species in intermittent streams, and highlight the importance of understanding the role of dispersal in species conservation and habitat restoration.     

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.     

Groundwater controls on nutrient cycling in a Mojave desert stream

1. Groundwater fluxes of nitrogen and dissolved organic carbon (DOC) were investigated in Grape Vine Canyon Stream in the Mojave Desert focusing on the rate of inputs and the fate of groundwater-derived nutrients in the stream. Discharge rates from different ground waters were measured using an end-member mixing model coupled with injections of a conservative solute tracer into the stream channel.
2. In surface water, nitrate concentration averaged 1.13 mg N L^–1 and DOC concentration averaged 1.82 mg C L^–1.
3. Groundwater discharge into Grape Vine Canyon Stream was derived from three sources. Nitrate concentration varied among the three groundwater sources with mean concentrations of 0.56, 0.94 and 0.08 mg N L^–1. DOC, in contrast, did not vary among ground water sources, with an overall average concentration of 2.96 mg C L^–1.
4. In the surface stream, nitrate concentration was two-fold greater than the concentration predicted from groundwater input, indicating that in-stream processes generated nitrate. Stream DOC concentration was lower than predicted based upon groundwater input rate. The production of nitrate and loss of DOC suggest that DOC is lost through mineralisation of dissolved organic matter, possibly resulting in the mineralisation of dissolved organic nitrogen to ammonium and subsequent transformation to nitrate via nitrification. In further support of this hypothesised linkage, DOC loss explained 80–89% of the variance in nitrate production in Grape Vine Canyon Stream.     

The roles of environment and space in shaping stream diatom communities

Diatoms are widely used in stream quality assessment due to their response to the local environment. Diatoms are also influenced by many large-scale processes and so the diatom communities of boreal streams incorporate a strong spatial component at a regional level. What is not properly known yet is whether the variation in diatom communities between regions is larger than the variation in measured environmental variables. We studied the roles of environment and space in accounting for variability in stream diatom communities across four regions in Finland. According to canonical correspondence analysis, geographical coordinates, nutrient concentrations (total N and P), and water conductivity were the most important factors affecting variation in diatom community composition. Of physical factors, depth and current velocity were also significant. According to Mantel tests, both environmental and geographical distances were related to dissimilarity in diatom community composition. Analysis of Similarities indicated that the regional differences in diatom community composition were larger than the regional differences in environmental variables. We also found many indicator species confined to certain regions. Our results suggest that the four study regions differ in their diatom species composition more than in their environmental features and that diatoms are structured not only by the local environment but also by large-scale processes, possibly related to history, climate and dispersal. These results imply that, while diatom species composition reflects well the environmental differences between regions, future bioassessments would benefit from regional stratification. Otherwise, relationships with environmental variables may be masked by trans-regional differences in species pools caused by the large-scale processes.     

Utilization of autochthonous macroinvertebrate drift by a pool fish community in a woodland stream

Qualitative and quantitative drift data were collected simultaneously above and below a pool both before and after the pool was heavily electrofished. These data revealed no significant difference between drift organism densities (#/m³) above or below the pool before or after fish collection. Qualitative and quantitative fish gut analyses suggest that the sunfishes Lepomis megalotis, L. macrochirus and L. cyanellus may be feeding on drifting invertebrates as these organisms comprised 58%, 37% and 35% gut volume, respectively. A design for a long-term drift net apparatus is presented.     

Physico-chemistry and aquatic insects of a glacier-fed and a spring-fed alpine stream

1. Physico-chemical conditions and benthic macroinvertebrates were studied in two adjacent alpine streams in the Tyrolean Alps, Austria, for 2 years, and aquatic insect emergence was recorded for 1 year.
2. In the spring-fed system, maximum discharge and increased concentrations of suspended solids, nitrate and particulate phosphorus occurred during snowmelt in June. In the glacier-fed stream, high discharge and strong diel fluctuations in flow and concentrations of suspended solids created a harsh and unstable environment during summer. Glacial ablation, variation in groundwater inflow, and water inputs from tributaries draining calcareous rocks caused water chemistry to vary both seasonally and longitudinally in glacier-fed Rotmoosache.
3. A total of 126 aquatic or semi-aquatic invertebrate taxa were collected, 94 of which were found in the glacier-fed stream and 120 in the spring-fed stream. Chironomid abundance was 2–8 times and taxa richness 2–3 times lower in the glacier-fed stream than in the spring-fed stream, as was the number of chironomid taxa (72 versus 93 total).
4. These results broadly support the conceptual model by Milner & Petts (1994) concerning glacier-fed stream systems. However, single samples and seasonal means showed relatively high invertebrate abundance and richness, especially during winter, indicating a considerable degree of spatial and temporal variability.
5. We suggest that the seasonal shifts from harsh environmental conditions in summer to less severe conditions in autumn and a rather constant environment in winter are an important factor affecting larval development, life-history patterns and the maintenance of relatively high levels of diversity and productivity in glacier-fed streams.     

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.     

Interspecific competition between Sitophilus zeamais and Sitotroga cerealella in a patchy environment

The maize weevil, Sitophilus zeamais (Motchulsky) (Coleoptera: Curculionidae), and the Angoumois grain moth, Sitotroga cerealella (Olivier) (Lepidoptera: Gelechiidae), are important insect pests of stored products. The coexistence in nature of the two species on maize has been difficult to explain from laboratory set‐ups, as the moth is inevitably eliminated by S. zeamais. However, early laboratory experiments ignored the spatial dimension, while several studies have lately revealed that two competing species may coexist in a spatially divided environment even though one of the species is competitively superior in a simpler universe. Two hypotheses for coexistence were examined by studying the interactions in a system where the resources were spatially divided. It was found that spatial subdivision in the current experiment was insufficient to ensure coexistence between the two species, so neither of the hypotheses could be confirmed. A simple model describing the population dynamics of S. zeamais was fitted to the data. Its parameters provide important information about how resource depletion influences the per capita growth rate. Finally, the study indicates that the mechanism responsible for S. zeamais's rapid elimination of S. cerealella is the superiority of the former species to colonize and monopolize new patches. The design of this study may serve as a template for further laboratory experiments revealing the effect of a spatially divided environment on competitive interactions. However, the present study also indicates that choosing the appropriate spatial scale may be critical for relating the results to natural environments.     

Hierarchy,spatial configuration,and nutrient cycling in a desert stream

Abstract Recent studies of nutrient cycling in Sycamore Creek in Arizona, USA, suggest that a thorough understanding requires a spatially explicit, hierarchical approach. Physical configuration determines the path that water follows as it moves downstream. Water follows flowpaths through surface stream components, the hyporheic zone beneath the surface stream, and the parafluvial (sand bar) zone. Characteristic biogeochemical processes in these subsystems alter nitrogen (N) species in transport, in part as a function of available concentrations of N species. At several hierarchical levels, substrate materials are an important determinant of nitrogen dynamics in desert streams. Sand is present in bars of variable size and shape, each of which can be considered a unit, interacting with the surface stream. Groups of these stream-sandbar units form a higher level, the reach. At the next higher scale, sandy reaches (runs) alternate with riffles. Where flowpaths converge, rates of N transformation are high and, as a result, change in concentration is a non-linear function of flowpath length. Disturbance by flash floods alters sandbar configuration. Between floods, the interaction of subsurface and surface flowpaths shapes configuration in each, thus a self-organizing element of spatial structure exists. Sandy runs are dominated by subsurface processes and are likely to be net nitrifiers while riffles are dominated by surface flow and are nitrogen fixers. Whether a stream ecosystem retains nitrogen, or transports it to downstream recipient systems, or is a net emitter of gaseous forms of N, depends upon the dynamics of a spatial mosaic of interacting elements. An understanding of the net effect of this mosaic requires a spatially explicit, hierarchical, multi-scale approach.