Predator impacts on stream benthic prey

The impact that predators have on benthic, macroinvertebrate prey density in streams is unclear. While some studies show a strong effect of predators on prey density, others show little or no effect. Two factors appear to influence the detection of predator impact on prey density in streams. First, many field studies have small sample sizes and thus might be unable to detect treatment effects. Second, streams contain two broad classes of predators, invertebrates and vertebrates, which might have different impacts on prey density for a variety of reasons, including availability of refuge for prey and prey emigration responses to the two types of predators. In addition, predatory vertebrates have more complex prey communities than predatory invertebrates; this complexity might reduce the impact that predatory vertebrates have on prey because of indirect effects. I conducted a meta-analysis on the results of field studies that manipulate predator density in enclosures to determine (1) if predators have a significant impact on benthic prey density in streams, (2) if the impacts that predatory invertebrates and vertebrates have differ, and (3) if predatory vertebrates have different impacts on predatory prey versus herbivorous prey. The results of the meta-analysis suggest that on average predators have a significant negative effect on prey density, predatory invertebrates have a significantly stronger impact than predatory vertebrates, and predatory vertebrates do not differ in their impact on predatory versus herbivorous invertebrate prey. Three methodological variables (mesh size of enclosures, size of enclosures, and experimental duration) were examined to determine if cross correlations exist that may explain the differences in impact between predatory invertebrates and vertebrates. No correlation exists between mesh size and predator impact. Over all predators, no correlation exists between experimental duration and predator impact; however, within predatory invertebrates a correlation does exist between these variables. Also, a correlation was found between enclosure size and predator impact. This correlation potentially explains the difference in impact between predatory invertebrates and predatory vertebrates. Results of the meta-analysis suggest two important areas for future research: (1) manipulate both types of predators within the same system, and (2) examine their impacts on the same spatial scale.     

Microdistribution of benthic invertebrates in a rocky mountain (U.S.A.) stream

A study of the benthic invertebrate community inhabiting a small, foothill trout stream in the Rocky Mountains of Idaho was conducted over a two-year period. Monthly Hess samples and short-term experiments using substratum-filled trays were used to describe the spatial dispersion of the benthos and to examine the response of invertebrate populations to substratum and current. A method was devised for measuring available surface area which involved coating individual stones with latex and measuring the area of the print resulting from inking the impression left on the latex mold.The dispersion of all populations was clumped throughout the year. Alteration of the cross-sectional pattern of current velocity and stream bed composition changed the pattern of distribution but not the extent of clumping. Collections made in areas of depositing and eroding substrata revealed a more diverse fauna in the latter. Most groups of organisms found in the riffle were scarcer in the pools or absent from them. The pool fauna contained no important additions over those found in the riffles.After a year's study of invertebrate populations in an otherwise undisturbed riffle, the substratum was altered and the flow made more uniform; an increase in the abundance of most of the benthic invertebrates followed. No single factor was responsible for the increase, but the change in substratum size and degree of compaction accounted for most of the change. Interpretation of the results was aided by findings from experiments using substratum-filled trays.Two series of stream experiments using the trays were conducted: one to test the relative importance of current and substratum and the other to test the effect of particle size on the distribution of the benthic fauna. In the first series, placement of trays of stones in a pool resulted in an increase in numbers of some but not all of the invertebrates over numbers usually occurring in the pool. Trays filled with stones and placed in a riffle supported fewer animals than found on the adjacent stream bed but more than in the pool. Variations are attributed to differences in current velocity and amounts of imported organic and inorganic debris. Three different relationships of population numbers to current velocity were found for different members of the community (direct, indirect, and parabolic) over the range of 10 to 60 cm/sec. The second series of experiments consisted of two sets of trays filled with stones of medium or large pebbles, respectively. Nine taxa, as well as all of the combined taxa, showed a preference for trays of small stones over the natural stream bed. A few taxa were noticeably more abundant on the small substratum than on the large but most of the fauna showed only slight increases in numbers or remained constant on the two substrata. Only three taxa showed a direct relation of numbers to total surface area presented by the stones.Number and kinds of organisms found in trays filled with a uniform size of substratum did not correspond to those taken in Hess samples from the natural stream bed. This has important implications in terms of currently recommended pollution monitoring techniques. However, it is suggested that if the substratum composition of the trays more nearly matched that of the stream, the correspondence would be much better. The results of the present study also throw considerable doubt on the adequacy of generalizations derived from earlier studies of responses to substratum size and suggest several reasons for reevaluating current ideas regarding the influence of substratum on invertebrate distribution.     

Effects of salinity gradients on benthic invertebrate and diatom communities in a German lowland river

The Lippe is a lowland river located in the Western part of Germany and has been heavily impacted by coal mining activities ever since. Although mining activities significantly decreased during the last decades, the associated discharge of salt-enriched mine water into the river still poses a persistent threat to the local benthic invertebrate and diatom communities. To analyze the effect of salt pollution on invertebrate and diatom species, biological and chemical data were compiled for this study from a publicly available database. Changes in the community composition due to increased salt concentrations were explored by Non-Metrical-Multidimensional Scaling. Indicator species and salinity thresholds for single species and communities were identified using the method TITAN (Threshold Indicator Taxa Analysis). The method is an analytical approach to detect changes in frequency and abundance of species along an environmental gradient by combining the methods of change point analysis (nCPA) and indicator species analysis. The obtained salinity preferences and individual and community thresholds were compared to the literature and existing salinity classifications. For both diatoms and benthic invertebrates, Non-Metrical-Multidimensional Scaling showed a clear split between samples of high and low salinities. Significant salinity thresholds were determined for 50 invertebrate and 58 diatom species of which 23 respectively 18 species were described as ‘reliable’ indicators according to the specifications given by Baker and King (2010). A majority of salt-tolerant indicator organisms were invasive species. For both organism groups, major changes in community composition were detected at a conductivity value exceeding 900 μS/cm. A reduction of the average salinity to below this threshold may have positive effects on the overall species richness and the persistence of sensitive taxa in the river Lippe. Individual and community thresholds may however be data-dependent to a certain degree and subjected to fluctuations considering the potential interdependencies between salinity and additional physico-chemical and environmental parameters (e.g. water temperature, lime content).     

Distribution and seasonal abundance of benthic macroinvertebrates in a subtropical Florida lake

We studied the distribution and seasonal abundance of benthic macroinvertebrates from July 1975 through September 1976 in a hypereutrophic lake in subtropical Florida. The benthic community was comprised principally of oligochaetes (56.1%), chironomids (37.1%), and chaoborids (5.7%). Numbers of taxa and mean densities correlated negatively with depth and positively with mean grain size of the substratum and dissolved oxygen concentration at the mud-water interface. Seasonal abundances and life history information obtained for the predominant species of Chironomidae (Polypedilum halterale, Glyptotendipes paripes, Chironomus crassicaudatus, Cryptochironomus fulvus, C. blarina, Cladotanytarsus sp., Procladius culiciformis, and Coelotanypus concinnus) indicated that all of these species are multivoltine with rapid generation times. Larval lengths of life at summer temperatures, 27–31 °C, ranged from 14–22 days indicating that sampling in subtropical lakes should be at short intervals (approximately 3 days) if the life cycles, ecology, and function of the components of the benthic community is to be understood.     

Diversity as a measure of benthic macroinvertebrate community response to water pollution

The assumption that water pollution causes a depression in the diversity of benthic macroinvertebrates as measured by the Shannon index and similar diversity indices is questioned. An interpretation of the community response of benthic macroinvertebrates to pollution in the Millers River, Massachusetts is developed from species presence-absence and abundance data in conjunction with published information on the species' environmental tolerances as compared to chemical water quality data. This interpretation is compared with one derived solely from diversity index values. The interpretations are quite different; the differences may be attributed to other environmental factors such as impoundments and flow reductions which influence the fauna and thus the diversity index value, but which are not related to pollution. In addition, several intrinsic features of the diversity indices increase their bias.Contribution No. 47 of the Massachusetts Cooperative Fishery Unit jointly supported by the U.S. Bureau of Sport Fisheries and Wildlife, the Massachusetts Division of Fisheries and Game, the Massachusetts Division of Marine Fisheries, and the University of Massachusetts.     

Primary producers in a Pampean stream: temporal variation and structuring role

Low current velocities, high nutrient levels, the lack of riparian forest vegetation, and the development of dense and rich macrophyte communities characterize Pampean streams. The objective of this study was to describe the main physical, chemical, and biological characteristics of a headwater Pampean stream as well as to analyze the role of macrophytes and phytobenthos. The study was conducted in a stream considered to be not much disturbed by human activities. Samples of water and organisms (macrophytes, benthic algae and invertebrates) were taken monthly for 14 months in two sampling stations, in fast flow and slow flow sites. Macrophyte biomass and diversity increased in spring and summer, and they decreased in autumn, when the plant community was greatly affected by an important flood. Phytobenthos biomass was lower in late summer, possibly due to the establishment of a dense cover of the floating macrophyte Lemna gibba L. Density of amphipods and gastropods greatly increases in spring and summer, jointly with the macrophyte development. Analysis of correlation showed that current velocity is the most important factor influencing macrophyte biomass and phytobenthos structure, while depth, nutrients, and herbivores are linked factors. Pampean streams could be considered systems dynamically fragile, because habitat heterogeneity is generated by aquatic vegetation, a substratum that varies along time.     

Development of a multimetric approach to assess perturbation of benthic macrofauna in Zostera noltii beds

Biotic indices based on soft-bottom macrozoobenthic communities are currently used throughout Europe to assess the ecological quality of coastal and transitional water bodies according to the European Water Framework Directive. However, the performance of the currently available biotic indices still has to be tested against a variety of different impact sources. In particular, physical perturbations have received much less attention than other kind of disturbances. This study consisted in testing the capacity of currently available uni- (BOPA, AMBI and BENTIX) and multivariate (M-AMBI) Biotic Indices to assess the ecological impact of the destruction of a Zostera noltii seagrass bed in Arcachon Bay (France) following sediment deposits. Changes of habitat after this physical perturbation were hardly assessed by any of these Biotic Indices whereas analysis of the benthic community showed drastic changes of structure following the perturbation and no recovery after 15 months. This study demonstrates that these Biotic Indices must be integrated into a multimetric approach which describes better the biological integrity of the benthic community by including a complementary set of metrics. A new multimetric approach, named MISS (Macrobenthic Index of Sheltered Systems) is proposed. MISS correctly highlighted the destruction of the seagrass beds, by using 16 metrics describing the biological integrity of the macrofauna.     

Quantitative hydrological preferences of benthic stream invertebrates in Germany

Current knowledge regarding the flow preferences of benthic stream invertebrates is mostly based on qualitative data or expert knowledge and literature analysis. These established flow preferences are difficult to use in predictions of the effects of global change on aquatic biota. To complement the existing categories, we performed a large-scale analysis on the distribution of stream invertebrates at stream monitoring sites in order to determine their responses to various hydrological conditions.We used 325 invertebrate surveys from environmental agencies at 238 sites paired to 217 gauges across Germany covering a broad range of hydrological conditions. Based on these data, we modelled the respective probabilities of occurrences for 120 benthic invertebrate taxa within this hydrological range using hierarchical logistic regression models.Our analyses revealed that more than one-third of the taxa (18–40%) can be considered as ubiquitous and having a broad hydrological tolerance. Furthermore, 22–41% of the taxa responded to specific ranges of flow conditions with detectable optima. “Duration high flow event” represented the flow parameter that correlated best with the abundance of individual taxa, followed by “rate of change average event”, with 41 and 38% of the taxa showing a peak in their probability of occurrence at specific ranges of these metrics, respectively. The habitat suitability for these taxa may be potentially affected by global change-induced hydrological changes.Quantified hydrological traits of individual taxa might therefore support stream management and enable the prediction of taxa responses to flow alteration. The hydrological traits of stream benthic invertebrates may be used in forecasting studies in central Europe, and the methods used in this study are suitable for application in other regions with different flow regimes.     

Fish distribution and benthic invertebrate biomass relative to depth in an Ontario lake

Synopsis A survey of fish distribution relative to depth in Lake Opinicon, Ontario, using the strip count method showed 80–90% of the biomass to be concentrated along the lake margins at a depth of up to 2.5 m. This figure applied throughout the summer, and to both day and night. Invertebrate diversity and biomass was also biassed towards the margins but slightly less so (mean summer figure 68% of biomass at depth of 2.5 m or less) for the segment of the lake studied. The central parts of the lake have good populations of 1–2 cm Chironomus spp. not predated by fish.There is a close link between the distribution of the specific prey organisms of fish species and the fish themselves. In their predominantly marginal distribution both are concentrated into the area of maximum productivity.     

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

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

Species aggregation: the influence of detritus in a benthic invertebrate community

Species aggregation with detritus was examined in a zoobenthic stream community. Detritus was quantified (dry weight in particle size categories) in contiguously arranged quadrats along four transects within a 40m stream section. Total organism density was found to be positively related to increasing detrital biomass (dry weight) in all transects, although the magnitude of this association was influenced by the detrital particle size distribution in each quadrat. Eliminating certain particle size categories of detritus resulted in substantial changes in the magnitude of association. For example, in one transect, total organism density and total detrital biomass showed no association. However, when one particle size category (greater than 10 mm) was removed a highly significant relationship was found. Total detrital biomass was found to be a useful predictor of organism distribution, however, inordinately high detrital biomass and variance within larger particle size categories can mask a relationship. This result suggests that the use of discrete size categories is necessary when considering species/ detrital relationships.Significant associations were found between the density of particular species and total biomass of detritus. The magnitude, as well as significance, of these associations were found to vary when correlations were calculated with restricted particle size categories of detritus. Species associations were often not consistent between transects, possibly due to local characteristics of detritus or overriding environmental factors. Current velocity was found to be unimportant in determining the detrital biomass held in the substrate. Likewise, few associations were found between current velocity and organism density in three out of four transects. Results of this study suggest that habitat selection by some taxa is influenced by local variation in detrital biomass, as well as detrital particle size composition.     

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

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

Drift and benthic invertebrate responses to stepwise and abrupt increases in non-scouring flow

We conducted two experiments to assess drift and benthic invertebrate responses to stepwise and abrupt changes in non-scouring flow in gravel-bed experimental streams. Intuitively, a stepwise flow increase should allow aquatic invertebrates more time to seek refuges than would an abrupt increase. We hypothesized that abrupt flow increases would result in larger increases in taxon richness and in the number of invertebrates in the drift, and a larger decrease in benthic density than would stepwise flow increases. Two kinds of drift response to flow increases were observed in the stepwise experiment: (1) no response (e.g. Caenissp. [Ephemeroptera] and Sphaeriidae [Pelecypoda]); (2) threshold response of some aquatic (e.g. Crangonyx pseudogracilis[Amphipoda]) and semiaquatic (e.g. Ormosiasp. [Diptera]) taxa. Drift richness and drift density in both experiments declined after reaching a peak. The peak was reached almost immediately in the abrupt treatment and later in the stepwise treatment. Maximum richness of taxa and taxon composition in the drift were similar in both experiments. Despite significant increases in drift, stepwise and abrupt increases in flow did not have a significant effect on benthic density. However, relative to reference streams, the percentage of total benthic invertebrates in the drift increased 10× in the stepwise experiment and 33× in the abrupt experiment. These non-scouring increases in flow were non-trivial. Our results suggest that several high flow events of the same magnitude (i.e. 2.5–3.0 fold increases) may cause considerable losses of benthic populations to the drift. The rate of increases in flow appears to be important: abrupt increases in flow had a stronger effect on invertebrate drift than did stepwise increases.     

Habitat scale and biodiversity: influence of catchment, stream reach and bedform scales on local invertebrate diversity

Although many studies have investigated the influence of environmental patterns on local stream invertebrate diversity, there has been little consistency in reported relationships between diversity and particular environmental variables. Here we test the hypothesis that local stream invertebrate diversity is determined by a combination of factors occurring at multiple spatial scales. We developed predictive models relating invertebrate diversity (species richness and equitability) to environmental variables collected at various spatial scales (bedform, reach and catchment, respectively) using data from 97 sampling sites dispersed throughout the Taieri River drainage in New Zealand. Models based on an individual scale of perception (bedform, reach or catchment) were not able to match predictions to observations (r < 0.26, P > 0.01, between observed and predicted equitability and species richness). In contrast, models incorporating all three scales simultaneously were highly significant (P < 0.01; r = 0.55 and 0.64, between observed and predicted equitability and species richness, respectively). The most influential variables for both richness and equitability were median particle size at the bedform scale, adjacent land use at the reach scale, and relief ratio at the catchment scale. Our findings suggest that patterns observed in local assemblages are not determined solely by local mechanisms acting within assemblages, but also result from processes operating at larger spatial scales. The integration of different spatial scales may be the key to increasing model predictability and our understanding of the factors that determine local biodiversity.     

Seasonal variation in benthic invertebrate recolonization of small-scale disturbances in a subtropical Florida lake

Approximately 11.5% of the littoral zone of a hypereutrophic Florida lake is disturbed by the construction of spawning beds by the cichlid fish, Sarotherodon aurea, during a single spawning season. Simulations of these beds were created during three seasons of the year to test for variation in recolonization rates and patterns in the benthic macroinvertebrate community.The seasonal variation in densities of benthic invertebrate populations suggests a direct relationship between reproductive activity and recolonization rate. Recolonization rates of the numerically dominant species, Polypedilum halterale (Diptera: Chironomidae), were much more rapid in the spring and summer than in the winter. In contrast, Hyalella azteca and Glyptotendipes paripes attained peak population densities during the winter season when Polypedilum was present in low densities. These organisms invaded the disturbed area in unusually high abundances and later declined to background levels. The final result was a winter population with densities comparable to the control (undisturbed) area, but the species composition was different.Similarity between disturbed and control communities during the winter season only reached 67% a month after disturbance, while communities during the warmer months attained nearly 90% similarity in less than 15 days. This lack of similarity during the winter indicates that disturbance, at the appropriate time, may play a role in community organization.     

Habitat effects on invertebrate drift in a small trout stream: implications for prey availability to drift-feeding fish

In this study, we focused on the drivers of micro- and mesohabitat variation of drift in a small trout stream with the goal of understanding the factors that influence the abundance of prey for drift-feeding fish. We hypothesized that there would be a positive relationship between velocity and drift abundance (biomass concentration, mg/m³) across multiple spatial scales, and compared seasonal variation in abundance of drifting terrestrial and aquatic invertebrates in habitats that represent the fundamental constituents of stream channels (pools, glides, runs, and riffles). We also examined how drift abundance varied spatially within the water column. We found no relationship between drift concentration and velocity at the microhabitat scale within individual pools or riffles, suggesting that turbulence and short distances between high- and low-velocity microhabitats minimize changes in drift concentration through settlement in slower velocity microhabitats. There were also minimal differences in summer low-flow drift abundance at the mesohabitat scale, although drift concentration was highest in riffle habitats. Similarly, there was no differentiation of drifting invertebrate community structure among summer samples collected from pools, glides, runs, and riffles. Drift concentration was significantly higher in winter than in summer, and variation in drift within individual mesohabitat types (e.g., pools or riffles) was lower during winter high flows. As expected, summer surface samples also had a significantly higher proportion of terrestrial invertebrates and higher overall biomass than samples collected from within the water column. Our results suggest that turbulence and the short length of different habitat types in small streams tend to homogenize drift concentration, and that spatial variation in drift concentrations may be affected as much by fish predation as by entrainment rates from the benthos. Handling editor: Robert Bailey     

Scour and fill patterns in a New Zealand stream and potential implications for invertebrate refugia

1. The hyporheic zone has long been regarded as a potential refugium for lotic invertebrates during disturbance. However, there have been few attempts to quantify the stability of this habitat during high flow events. In a New Zealand stream with an unstable bed, the present authors monitored spatial patterns of scour and fill in a riffle in a wide flood plain and at two sites in a constrained reach: a pool-riffle with bedrock outcrops and a plane-bed (a bedform characterized by long stretches of planar stream bed). 2. At each 20-m site, 100 scour chains were installed in a systematic grid with about 1 m between chains. Scour was measured by comparing the length of chain exposed before and after a high flow event, whereas filling depth was equivalent to the thickness of the sediment deposited on top of the chains during the event. For each chain, the present authors noted dominant particle size and degree of packing of the surrounding bed, water depth and presence or absence of large stones upstream. Chains were re-located after four smaller spates, one intermediate event and one large flood. 3. Most events caused a complex mosaic of bed patches which experienced scour, fill or remained undisturbed. These patterns, which were mostly site- and event-specific, were often significantly influenced by the longitudinal or lateral position of the chains in the spatial grids. 4. The cumulative effect of the six high flow events differed substantially between sites. The first site experienced predominantly scour, the second both scour and fill, and the third almost exclusively fill. These differences were partly explained by channel geomorphology. The bedrock outcrops at the constrained pool-riffle site forced the flow at high discharge, causing deep scour in these areas, whereas a backwater effect at the third site reduced near-bottom shear stress during larger events and led to sediment deposition. 5. Except for a single event at the second site, scour affected mainly the uppermost 10–15 cm of the stream bed. Therefore, almost the entire hyporheic zone below this depth would have been available as refugium for invertebrates, in addition to the often consider-able number of bed patches which remained undisturbed during the six high flow events. 6. Fill without earlier scour during the same high flow event was common at all sites. Most previous studies have assumed that lotic invertebrates are mainly affected by scour during high flow events, but the consequences of sediment deposition may be just as far reaching.     

The impact of vertebrate and invertebrate predators on a stream benthic community

I assessed the impact of both vertebrate and invertebrate predators on a lotic benthic community in a 1-month-long experiment, using enclosures containing cobble/gravel bottoms, with large-mesh netting that allowed invertebrates to drift freely. Brown trout (Salmo trutta) and leeches (Erpobdella octoculata) were used as predators and four treatments were tested: a predator-free control, leeches only, trout only, and leeches and trout together. A density of 26.7 leeches/m² (20 leeches/enclosure) and 1.3 trout/m² (one trout per enclosure) was stocked into the enclosures. The total biomass of invertebrate prey was significantly lower in the trout and trout plus leech treatments than in the leech and control treatments, which were due to strong negative effects of trout on Gammarus. On the individual prey taxon level, both trout and leeches affected the abundance of Asellus , Baetis and Ephemerella, whereas the abundance of Gammarus was only affected by trout, and the abundance of Orthocladiinae and Limnephilidae was only affected by leeches. In the treatment with trout and leeches together, the abundance of Ephemerella and Baetis was higher than when trout or leeches were alone, which was probably due to predator interactions. Leeches and trout had no effects on prey immigration but did affect per capita emigration rates. Both trout and leeches indirectly increased periphyton biomass in enclosures, probably due to their strong effects on grazers. Both trout and leeches were size-selective predators, with trout selecting large prey, and leeches selecting small prey. Size-selective predation by trout and leeches affected the size structure of five commonly consumed prey taxa. Trout produced prey populations of small sizes owing to consumption of large prey as well as increased emigration out of enclosures by these large prey. Leech predation produced prey assemblages of larger size owing to consumption and increased emigration of small prey. These results suggest that in lotic habits, predatory invertebrates can be as strong interactors as vertebrate predators. Received: 23 June 1997 / Accepted: 4 May 1998     

Nitrate retention in a sand plains stream and the importance of groundwater discharge

We measured net nitrate retention by mass balance in a 700-m upwelling reach of a third-order sand plains stream, Emmons Creek, from January 2007 to November 2008. Surface water and groundwater fluxes of nitrate were determined from continuous records of discharge and from nitrate concentrations based on weekly and biweekly sampling at three surface water stations and in 23 in-stream piezometers, respectively. Surface water nitrate concentration in Emmons Creek was relatively high (mean of 2.25 mg NO₃?CN l^?1) and exhibited strong seasonal variation. Net nitrate retention averaged 429 mg NO₃?CN m^?2 d^?1 and about 2% of nitrate inputs to the reach. Net nitrate retention was highest during the spring and autumn when groundwater discharge was elevated. Groundwater discharge explained 57?C65% of the variation in areal net nitrate retention. Specific discharge and groundwater nitrate concentration varied spatially. Weighting groundwater solute concentrations by specific discharge improved the water balance and resulted in higher estimates of nitrate retention. Our results suggest that groundwater inputs of nitrate can drive nitrate retention in streams with high groundwater discharge.