Recovery of three arctic stream reaches from experimental nutrient enrichment

1. Nutrient enrichment and resulting eutrophication is a widespread anthropogenic influence on freshwater ecosystems, but recovery from nutrient enrichment is poorly understood, especially in stream environments. We examined multi‐year patterns in community recovery from experimental low‐concentration nutrient enrichment (N + P or P only) in three reaches of two Arctic tundra streams (Kuparuk River and Oksrukuyik Creek) on the North Slope of Alaska (U.S.A.). 2. Rates of recovery varied among community components and depended on duration of enrichment (2–13 consecutive growing seasons). Biomass of epilithic algae returned to reference levels rapidly (within 2 years), regardless of nutrients added or enrichment duration. Aquatic bryophyte cover, which increased greatly in the Kuparuk River only after long‐term enrichment (8 years), took 8 years of recovery to approach reference levels, after storms had scoured most remnant moss in the recovering reach. 3. Multi‐year persistence of bryophytes in the Kuparuk River appeared to prevent recovery of insect populations that had either been positively (e.g. the mayfly Ephemerella, most chironomid midge taxa) or negatively (e.g. the tube‐building chironomid Orthocladius rivulorum) affected by this shift in dominant primary producer. These lags in recovery (of >3 years) were probably driven by the persistent effect of bryophytes on physical benthic habitat. 4. Summer growth rates of Arctic grayling (both adults and young‐of‐year) in Oksrukuyik Creek (fertilised for 6 years with no bryophyte colonisation), which were consistently increased by nutrient addition, returned to reference rates within 1–2 years. 5. Rates of recovery of these virtually pristine Arctic stream ecosystems from low‐level nutrient enrichment appeared to be controlled largely by duration of enrichment, mediated through physical habitat shifts caused by eventual bryophyte colonisation, and subsequent physical disturbance that removed bryophytes. Nutrient enrichment of oligotrophic Arctic stream ecosystems caused by climate change or local anthropogenic activity may have dramatic and persistent consequences if it results in the colonisation of long‐lived primary producers that alter physical habitat.     

THE TROPHIC ECOLOGY OF FRESHWATER GAMMARUS SPP. (CRUSTACEA:AMPHIPODA): PROBLEMS AND PERSPECTIVES CONCERNING THE FUNCTIONAL FEEDING GROUP CONCEPT

Gammarus spp. are widespread throughout a diverse range of freshwater habitats and can be the dominant part of many benthic macroinvertebrate assemblages, in terms of both numbers and/or biomass. Although the vast majority of studies have emphasized the herbivorous nature of Gammarus spp. and their ‘shredder’ functional feeding group (FFG) classification, we show that a far wider food base is exploited than has been previously acknowledged. This ‘plasticity’ as herbivore/predator is linked to the success of Gammarus spp. in persisting in and colonizing/invading disturbance-prone ecosystems. Intraguild predation and cannibalism are more common than previously realized. This behaviour appears to be a causal mechanism in many amphipod species replacements. Additionally, Gammarus spp. are major predators of other members of the macroinvertebrate community. Furthermore, while many studies have emphasized fish predation on Gammarus spp., we illustrate how this fish: amphipod, predator: prey interaction may be a two-way process, with Gammarus spp. themselves preying upon juvenile and wounded/trapped fish. We urge that a new realism be adopted towards the trophic ecology of Gammarus spp. and their role as predators and prey and that previously established FFG assumptions of both the food and the feeder be questioned critically.     

Effects of freshwater shrimp assemblages on benthic communities along an altitudinal gradient of a tropical island stream

1. In tropical island stream ecosystems freshwater shrimps are often the dominant macroconsumers and can play an important role in determining benthic community composition. However, most studies of the ecological role of shrimps are limited to high‐altitude shrimp‐dominated sites where other biota (fishes and snails) are absent or significantly less abundant than at lower altitudes. 2. We examined how effects of different shrimp assemblages on benthic communities changed along an altitudinal gradient in a tropical island stream in Puerto Rico. We used electroshocking and observations to quantify abundance and taxonomic composition of shrimp assemblages at three sites (300, 90 and 10 m a. s. l) along the Río Espíritu Santo. We also experimentally manipulated access of shrimps to the benthic environment simultaneously at each site using electric fences over a 35‐day period. 3. At the high‐altitude site, exclusion of shrimps (predominantly Atya spp. and Xiphocariselongata) resulted in significantly greater accrual of organic and inorganic material, chlorophyll a and algal biovolume. In the absence of shrimps, the algal community was dominated by filamentous green algae (Chlorophyta: Oedogonium and Rhizoclonium). Excluding shrimps did not affect total insect biomass but significantly increased sessile chironomids (Diptera: Chironomidae). We observed similar treatment effects at the mid‐altitude site where shrimps (primarily Macrobrachium spp. and X. elongata) occurred at lower densities. In contrast, at the low‐altitude site there were no treatment differences in organic and inorganic material, chlorophyll a, algal biovolume, algal assemblage composition and insects. 4. The lack of treatment differences at the low‐altitude site was probably because of very high densities of grazing snails (Thiaragranifera and Neritina spp.) which reduced organic and inorganic resources and obscured potential shrimp effects. 5. This study demonstrates that freshwater shrimps can play an important role in determining benthic community composition; however, their effects vary and appear to depend on the presence of other biota. This study suggests that loss of shrimps as a result of anthropogenic disturbances will have different effects on the stream community depending upon location along the altitude gradient.     

Parallels and contrasts between intermittently freezing and drying streams: From individual adaptations to biodiversity variation

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Assessing the functional importance of large‐bodied invertebrates in experimental headwater streams

Recent theoretical advances in food web ecology emphasize the importance of body size disparities among species for the structure, stability and functions of ecosystems. Experimental confirmations of the functional importance of large species, independent of their trophic position, are scarce. We specifically examine the multiple ecological roles of large invertebrates from two distinct trophic levels in headwater streams. We experimentally manipulated the presence of large predatory invertebrates (two Perlid stoneflies) or detritivores (a limnephilid caddisfly and a Pteronarcys stonefly) in a two‐by‐two design in stream channels open to immigration/emigration of smaller biota. We assessed treatment effects on the trophic structure of the benthic invertebrate community, dynamics of basal resources (benthic algae and leaf litter of cedar and alder), and stability of litter decomposition rates against an experimental pulse perturbation (fine sediment input). The presence of the large invertebrates was associated with a ten‐fold decrease in the biomass of invertebrate filterers whereas other trophic groups were unaffected by the large species. The biomass of benthic algae was lower and the rate of mass loss of alder litter was higher in channels lacking the large predators, thus revealing trophic cascades operating along both algal‐based and detritus‐based food chains. The large predators had no detectable effect on the decomposition of cedar whereas both cedar and alder disappeared faster in the presence of the large detritivores. Furthermore, the large predators and large detritivores interactively influenced the decomposition of the cedar–alder mixture through a litter diversity effect and the variability of the rate of alder decomposition after a pulse of fine sediment. Because the large invertebrates affected multiple ecosystem properties, and as their absence was not rapidly compensated for by small immigrant species, our findings support the notion that large species could be critically important in controlling ecosystem structure and functioning.     

Separating physical disturbance and nutrient enrichment caused by Pacific salmon in stream ecosystems

1. Pacific salmon (Oncorhynchus spp.) deliver marine‐derived nutrients to the streams in which they spawn and die, and these resource subsidies can increase the abundance of stream biota. In strong contrast, physical disturbance from salmon spawning activity can reduce the abundance of benthic organisms. Previous experimental designs have not established the relative effects of these two contrasting processes on stream organisms during a salmon run. 2. We combined manipulative and observational field studies to assess the degree of nutrient enrichment, physical disturbance, and the net effect of salmon on the abundance of benthic periphyton. Related salmon‐mediated processes were also evaluated for benthic macroinvertebrates. Mesh exclosures (2 × 2 m plots) prevented salmon from disturbing areas of the stream channel, which were compared with areas to which salmon had access. Sampling was conducted both before and during the late‐summer spawning run of pink (O. gorbushca) and chum (O. keta) salmon. 3. Streamwater nitrogen and phosphorus concentrations increased sharply with the onset of the salmon run, and highly significant positive relationships were observed between the numbers of salmon present in the stream and these dissolved nutrients. Before the salmon run, periphyton biomass (as chlorophyll a) and total macroinvertebrate abundance were very similar between control and exclosure plots. During the salmon run, exclosures departed substantially from controls, suggesting significant disturbance imparted on benthic biota. 4. Comparing exclosures before and during the salmon run enabled us to estimate the effects of salmon in the absence of direct salmon disturbance. This ‘nutrient enrichment potential’ was significant for periphyton biomass, as was a related index for macroinvertebrate abundance (although enhanced invertebrate drift into exclosures during the salmon run could also have been important). Interestingly, however, the net effect of salmon, evaluated by comparing control plots before and during the salmon run, was relatively modest for both periphyton and macroinvertebrates, suggesting that nutrient enrichment effects were largely offset by disturbance. 5. Our results illustrate the importance of isolating the specific mechanisms via which organisms affect ecosystems, and indicate that the relative magnitude of salmon nutrient enrichment and benthic disturbance determines the net effect that these ecologically important fish have on stream ecosystems.     

Calcium concentration in leaf litter affects the abundance and survival of crustaceans in streams draining warm–temperate forests

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Multiple‐stressor effects on stream invertebrates: a mesocosm experiment manipulating nutrients,fine sediment and flow velocity

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The dynamics of predation on Gammarus spp. (Crustacea: Amphipoda)

Gammarus spp. (Crustacea: Amphipoda) are widespread throughout a diverse range of marine, freshwater and estuarine/brackish habitats, often dominating benthic macroinvertebrate communities in terms of both numbers and/or biomass. Gammarus spp. are the dominant macroinvertebrate prey items of many fish, whether as a seasonal food source or a year-round staple. Selective predation by fish on Gammarus spp. is often linked to parasitism and the body size of the prey. Gammarus spp. populations are under increasing threat from both pollution and replacement/displacement by introduced species. Loss of populations and species invasions/replacements could have significant impacts on native predator species if the predator(s) cannot successfully adapt their feeding patterns to cope with non-indigenous Gammarus prey species. Despite this, many fish predation studies do not identify Gammarus prey to species level. This lack of precision could be important, as Gammarus spp. exhibit wide variations in physiochemical tolerances, habitat requirements, abilities to invade and susceptibility to replacement. Although rarely acknowledged, the impacts of nonpiscean predators (particularly macroinvertebrates) on Gammarus prey species may frequently be stronger than those exerted by fish. A major aim of this review is to ascertain the current importance of Gammarus as a prey species, such that the implications of changes in Gammarus spp. populations can be more accurately assessed by interested groups such as ecologists and fisheries managers. We also review the dynamics of Gammarus spp. as prey to a diverse array of mammals, birds, amphibians, insects, flatworms, other crustaceans such as crabs and crayfish and, perhaps most importantly, other Gammarus spp.     

Faecal pellets in streams: their binding, breakdown and utilization

1. Faecal pellets of Gammarus (shredders) and Simulium larvae (suspension feeders) are bound by exopolymers. Immediately after egestion, Gammarus pellets are covered by a peritrophic membrane that breaks up within hours, although pellets remain intact because of internal binding materials. 2. Although they expand soon after egestion, the faecal pellets of Gammarus and Simulium remain intact for more than 30 days. Their internal structure is altered and the main agents of this change are bacteria that have survived passage through the gut (and become bound within pellets). 3. When disrupted physically, freshly egested (1‐ to 2‐day old) Simulium faecal pellets break up into relatively large pieces whereas freshly egested Gammarus faecal pellets break apart into much smaller pieces. Disruption of 30‐day old Simulium faecal pellets results in similar sized pieces to those from freshly egested pellets, but disruption of 30‐day old Gammarus pellets produces pieces that are two orders of magnitude larger than those resulting from disruption of freshly egested pellets. 4. Faecal pellets of Gammarus and Simulium are eaten by stream invertebrates and are sites of microbial breakdown. Faecal pellets are a source of organic matter for benthic invertebrates, bacteria and, indirectly, for plants.     

Responses of benthic algal communities and their traits to experimental changes in fine sediments,nutrients and flow

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Variability in the settlement of non-indigenous species in benthic communities from an oceanic island

The introduction of non-indigenous species (NIS) in new environments represents a major threat for coastal ecosystems. A good understanding of the mechanisms and magnitude of the impact of NIS colonisation on native ecosystems is becoming increasingly crucial to develop mitigation measures and prevent new invasions. In this present study, we asked if distinct coastal benthic communities from an oceanic island can have different vulnerability to NIS colonisation process. First, PVC settlement plates were deployed for 1 year on the rocky shore of two different locations of Madeira Island (North versus South coast). Then, we implemented a mesocosm experiment where recruited plate communities were maintained under different levels of NIS propagule pressure in order to assess their vulnerability to NIS colonisation process. Results showed that NIS colonisation success was not influenced by the level of propagule pressure, but however, final colonisation patterns varied depending on the origin of the communities. This variability can be attributed to major structural differences between the preponderant species of each community and therefore to the biotic substrate they offer to colonisers. This study highlights how biotic features can alter the NIS colonisation process and importantly, shows that in an urbanisation context, the nature of the resident communities facing invasions risks needs to be closely assessed.     

Facilitation of Native Stream Fauna by an Invading Species? Experimental Investigations of the Interaction of the Snail, Potamopyrgus antipodarum (Hydrobiidae) with Native Benthic Fauna

Biological invasions are regarded as a major threat to native ecosystems, yet studies investigating the interactions of invaders with native biota under field conditions are rare. Whilst many invaders are present only in low densities, it is often the effects of high densities that are of particular concern. We manipulated densities of the invading New Zealand aquatic snail Potamopyrgus antipodarum within an Australian stream to test the relationships between relatively high and low densities of the invading snail and other benthic fauna. Two experiments were carried out: the first experiment investigated the relationship between P. antipodarum and benthic fauna colonising within a short period of six days, the second looked at the effects after six days of high P. antipodarum densities on resident benthic fauna. In both experiments, there was no evidence of a negative relationship between densities of P. antipodarum and native fauna. On the contrary, both experiments showed a positive relationship between P. antipodarum densities and those of some common native fauna. In the second experiment invader densities were positively correlated with total number of native taxa, as well as with total densities and the densities of common invertebrates. Coprophagy is suggested as a possible mechanism by which increase in P. antipodarum could facilitate increase in native fauna. The results of this work strongly suggest that the effects of an invader can include indirect effects on the trophic dynamics of an ecosystem.     

Changes in riverine benthic diatom community structure along a chloride gradient

The application of salt is the primary means of deicing roads and highways in colder regions of north-eastern North America. This has increased the chloride concentrations of many lake and stream ecosystems. While this salinization has been documented, less is known about how increased salinity alters benthic communities in downstream ecosystems. Natural thresholds, at which there are large scale changes in community composition, have not yet been established for many types of contaminants, including chloride. The diatom community, which is sensitive to small changes in the ambient environment, has the potential to be a strong indicator of salinization effects on stream ecosystems. In this study, we sampled diatom communities in 41 streams across a salinity gradient throughout south central Ontario, Canada. We sampled benthic diatom assemblages in early May following complete snowmelt, when stream water chloride concentrations ranged from 5 to 502 mg/L. Based on redundancy analysis, we confirmed a strong association between the species composition of the diatom community and water conductivity, a commonly used index of stream salinity. Taxa indicator threshold analyses (TITAN) indicated the community changed substantially at chloride concentrations greater than 35 mg/L. We also found that, an indicator taxa, Meridion circulare, was extremely sensitive to high concentrations of salt and negatively correlated with chloride. In a wide synoptic survey of streams of our region, we found that streams in most developed watersheds exceed tolerance thresholds for benthic diatom communities. This work suggests that current chloride concentrations in urban watersheds are greatly exceeding the benthic community thresholds, for which improved management and regulatory practices are needed. Salinization thus appears to be an important feature of urban streams and needs to be considered as an important ecological driver in future studies.     

Sediment destabilizing and stabilizing bio-engineers on tidal flats: cascading effects of experimental exclusion

Bioturbating lugworms (Arenicola marina) were excluded from 400 m2 plots of intertidal sand which initiated sequences of direct and indirect changes in the structure of the benthic community. The sessile, tube-building species Polydora cornuta and Lanice conchilega took advantage of the absence of lugworms and settled preferentially on lugworm exclusion plots. The protruding tubes provided attachment for an ephemeral development of algal tufts (Berkeleya colonies and Enteromorpha thalli) which in turn enhanced settlement of the juvenile drifting clams Mya arenaria and Macoma balthica. This causal chain of enhanced bivalve settlement in the presence of above-ground structures, like animal tubes and algae, on lugworm exclusion plots occurred in 2 years at different tidal zones with different tube builders, algae and juvenile clams. A significant response of L. conchilega in a year with relatively low lugworm abundances at the entire site suggests that not only the actual absence of large bioturbators was responsible for the establishment of tube-dwelling species, but also a cumulative change of the sediment in exclusion plots since the onset of the experiment. While the sediment on lugworm plots remained permeable, fine particles and organic matter accumulated at exclusion plots. It is suggested that these differences in sediment characteristics were the product of divergent benthic engineering by sediment destabilizing lugworms on control plots and sediment stabilizing species on exclusion plots. Cumulative changes of the sedimentary habitat and cascading effects in the benthic community may explain the persistence of patches that are dominated by either sediment stabilizing or destabilizing species in the assemblage mosaic of intertidal sediments.     

Stable isotope proxies for evaluating biodiversity in stream biota

Differences in δ¹³C and δ¹⁵N values in stream biota are caused by several environmental conditions. Variations in abundance, species richness and the assemblage structure of stream biota are also caused by several environmental conditions. Hence, abundance, species richness and the assemblage structure of stream biota are expected to be strongly correlated with the differences in value of stable isotopes. In this study, the gaps in δ¹³C and δ¹⁵N between periphyton and charr are discussed in terms of the abundance, genus richness, and assemblage of benthic invertebrates at each site. Gaps in δ¹³C between periphyton and charr were strongly correlated with some aspects of mountainous area and the genus richness of benthic invertebrates at each site. The gaps in δ¹⁵N between periphyton and charr were strongly correlated with the abundance and assemblage structure of benthic invertebrates at the location tested. The δ¹³C values of predators were correlated with some aspects of mountainous area and the assemblage structure of the benthic invertebrates. The δ¹⁵N values of predators were correlated with genus richness and the assemblage structure of the benthic invertebrates. These results suggest that the value gaps in δ¹³C and δ¹⁵N can be used to assess biodiversity and could provide indices for estimating the biodiversity in a stream.     

THE BENTHIC MACROINVERTEBRATE FAUNA OF A SOUTH AFRICAN MOUNTAIN STREAM AND ITS RESPONSE TO FIRE

Summary

The effects of a late-summer prescribed burn on the temperature and benthic macroinvertebrate fauna of a south-western Cape mountain stream were investigated over a period of 12 months. Temperature and discharge regimes appear well-defined and relatively predictable from year to year. As in other mediterranean-type ecosystems, seasonal changes in the structure of the invertebrate community and the relative abundance of different feeding groups appear to be associated primarily with changes in the physical environment. Distinctive summer and winter communities were identified, with chironomids dominating the fauna in summer and simuliids dominant in winter. Although the riparian vegetation was only slightly damaged by the fire, a heavy, aseasonal leaf-fall occurred shortly afterwards. The canopy remained sparse for approximately four months. Stream temperature in the post-burn year was not demonstrably affected by increased exposure to solar radiation, however, probably because the canopy remained open during the winter months. The fire appeared to have little effect on the invertebrate fauna. Apart from five rare elements of the biota, all species recorded in the pre-burn year were present in the post-burn year and in similar densities. It is concluded that the riparian vegetation is of major importance in maintaining the integrity of the stream environment.     

Short-term predator avoidance behavior by invasive and native amphipods in the Great Lakes

Understanding predator avoidance behavior by prey remains an important topic in community and invasion ecology. Recently, the Ponto-Caspian amphipod Echinogammarus ischnus (Stebbing 1898) was accidentally introduced into the Great Lakes. Since its introduction, it has displaced the native amphipod, Gammarus fasciatus (Say 1818), from several locations in the lower lakes. To assess whether behavioral differences in predator avoidance might be a causal mechanism increasing the success of the invasive amphipods, two experiments were conducted examining (1) native and invasive amphipod behavioral responses to five fish species with different foraging behaviors, and (2) amphipod responses to different densities of round gobies, a hyper-abundant benthic invertivore. Echinogammarus reduced its distance moved in the presence of all fish species tested, whereas Gammarus reduced its distance moved only after exposure to round gobies, black crappies, and rainbow darters. Both amphipod species increased the time spent motionless following exposure to round gobies, but not after encountering the scent of most of the remaining fish predators. The exception was that Echinogammarus also responded to black crappie scent whereas Gammarus did not. Although both amphipod species exhibited behavioral responses to many of the fish predators, the magnitude of their responses differed only after exposure to the brown bullhead. In the bullhead trials, Echinogammarus reduced its distance traveled significantly more than Gammarus. Both amphipod species increased their avoidance response to increasing goby density, however, the pattern of avoidance behavior was different. Invasive E. ischnus exhibited a consistently strong avoidance response to round gobies over the test duration. Native G. fasciatus initially avoided goby scent, but then either ceased their avoidance response or showed a hyper-avoidance response, depending on goby density. These results suggested (1) both species of amphipods were able to differentiate and react to a variety of fish predators, (2) invasive Echinogammarus amphipods avoided a larger range of fish predators than the native Gammarus, (3) increased avoidance behavior was associated with an increased density of fish, and (4) the avoidance response patterns of invasive Echinogammarus when faced with round goby predators might lead to increased predation on native Gammarus in habitats where they co-occur.     

Fish predation affects the structure of a benthic community

1. We conducted an experimental study of predation by benthivorous fish on a natural community of stream invertebrates using a reach‐scale approach. Over a 2‐year period (experimental phase), the benthic invertebrate community of a stretch containing two species of benthivorous fish was compared with a fishless stretch. Thereafter, all fish were removed and benthic community structure was analysed again to account for natural differences between the two stretches (reference phase). 2. Benthivorous fish at the moderate densities investigated did not affect total benthic biomass or density, but did alter species composition. In addition, the fish effect differed between pool and riffle habitats, with larger effects in the pools indicating a habitat‐specific predation effect. In the reference phase, when all fish were removed from the stream, the difference between the two stretches was reduced. 3. The benthivorous fish reduced the densities of four taxa (Pisidium sp., Dugesia gonocephala, Gammarus pulex, Limoniidae), representing 29% of total biomass. It is possible that density reductions of other species were masked by prey migration despite the relatively large spatial scale. Indeed, higher drift activity in the upstream fishless stretch could have increased the density of Baetis rhodani in the fish stretch, as indicated by the results of a drift model. 4. Our results provide insights into stream food web ecology because fish predation showed effects even in a natural system where habitat complexity was high, environmental factors were highly variable and many predator and prey species interacted and because benthivorous fish were the focus, whereas the majority of previous predation experiments in streams have used drift‐feeding trout.     

Uncoupling strong predator–prey interactions in streams: the role of marginal macrophytes

Predation refugia can facilitate the coexistence of predators and prey within an ecosystem by weakening trophic links between the two. The marginal macrophytes of shallow lakes are used facultatively by zooplankton to escape fish predation, leading to the stabilisation of lentic food webs. Little is known about such a role for the marginal macrophytes of lotic systems. In this paper, we examine whether the marginal macrophytes of chalk streams help buffer the interaction between the freshwater shrimp, Gammarus pulex and a benthic fish predator, Cottus gobio, both characteristically abundant in such systems. Quantitative surveys were taken of Gammarus and bullheads in winter and summer in a southern English chalk stream. These indicated that Gammarus seasonally switched their habitat preference, from marginal macrophytes in summer, to midchannel habitats in winter. Bullheads exhibited an opposite trend, preferring midchannel habitats in summer and all habitats, particularly marginal macrophytes, in winter. Large Gammarus and precopula pairs were found almost exclusively in the margins in summer. This spatial separation between Gammarus and bullheads in summer, though not in winter, was reflected in bullhead diets, as determined by gut analysis. In field manipulation experiments, bullhead presence was the strongest factor explaining the between‐habitat distribution of Gammarus in both summer and winter, indicating that the habitat shift of Gammarus was driven by the distribution of fish. Other abundant invertebrate taxa, including Asellus, mayflies and chironomids, exhibited little avoidance of habitat patches of high bullhead density. We conclude that marginal macrophytes in chalk streams can potentially facilitate the coexistence of high densities of both Gammarus and bullheads by spatially separating predator and prey in summer. They may further allow large populations of Gammarus to persist in the presence of high bullhead density at stretch‐wide spatial scales, by reducing the predation by bullheads of large breeding and newly born individuals of Gammarus in summer.