Patchy bed disturbance and fish predation independently influence the distribution of stream invertebrates and algae

1. The identification of factors determining the patchy distribution of organisms in space and time is a central concern of ecology. Predation and abiotic disturbance are both well-known drivers of this patchiness, but their interplay is still poorly understood, especially for communities dominated by mobile organisms in frequently disturbed ecosystems. 2. We investigated the separate and interactive influences of bed disturbance by floods and predation by fish on the benthic community in a flood-prone stream. Electric fields excluded fish predators from half of 48 stream bed patches (area 0·49 m(2) ) with contrasting disturbance treatments. Three types of bed disturbance were created by either scouring or filling patches to a depth of 15-20 cm or by leaving the patches undisturbed, thus mimicking the mosaic of scour and fill caused by a moderate flood. Benthic invertebrates and algae were sampled repeatedly until 57 days after the disturbance. 3. Disturbance influenced all ten investigated biological response variables, whereas predation affected four variables. Averaged across time, invertebrate taxon richness and total abundance were highest in stable patches. Algal biomass and densities of five of the seven most common invertebrate taxa (most of which were highly mobile) were higher in fill than in scour patches, whereas two taxa were more abundant in scour and stable than in fill patches. Furthermore, two common invertebrate grazers were more abundant and algal biomass tended to be reduced in fish exclusion patches, suggesting a patch-scale trophic cascade from fish to algae. 4. Our results highlight the importance of patchy physical disturbance for the microdistribution of mobile stream organisms and indicate a notable, but less prevalent, influence of fish predation at the patch scale in this frequently disturbed environment. Disturbance and predation treatments interacted only once, suggesting that the observed predation effects were largely independent of local bed disturbance patterns.     

Are impacts of an exotic predator on a stream food web influenced by disturbance history?

Predatory species have been introduced to habitats spanning a wide range of environmental conditions. To better understand the consequences of predation in natural communities we need to examine how variations in abiotic factors modify the influence of predation. The effects of introduced predators may vary amongst habitats if natural disturbance affects the abundance and taxonomic composition of consumers and their resources, or the predator alters recolonisation after disturbance. We tested whether a bed-moving disturbance altered subsequent interactions involving native and introduced predatory fish, invertebrate grazers and algae in experimental channels within a New Zealand stream. Disturbance reduced the abundance of invertebrates by 84%, and induced mortality of Conoesucidae caddisflies. However, the relative abundance of taxa changed little immediately following the disturbance. Invertebrate communities recovered following disturbance in fishless channels and those with native galaxiids (Galaxias vulgaris), and were almost indistinguishable from undisturbed fishless controls after 2 weeks. Invertebrate abundance declined and algal abundance increased in channels with exotic brown trout (Salmo trutta) and their effect was strongest in previously disturbed channels. However, predators and disturbance only had interactive effects on grazer emigration rates. Trout affected grazers through direct consumption (e.g. Conoesucidae caddisflies), and induced higher emigration rates of grazers from channels via drift (e.g. the mayfly Deleatidium). The effects of predatory trout and galaxiids combined differed in disturbed and stable channels. The observed combined effects of predatory trout and galaxiids on invertebrate grazers were lower than expected in stable channels partly due to low emigration rates of Conoesucidae, whereas emigration of grazers was higher than expected in the disturbed channels. The biomass of algae was higher than expected in disturbed channels with both predators. Collectively, our results indicate that predator substitutability and the non-lethal effects of introduced predators varied depending on disturbance history, but their effects on the biomass of grazers and algae did not.     

The influence of introduced trout on the benthic communities of paired headwater streams in the Sierra Nevada of California

1. Non‐native trout have been stocked in streams and lakes worldwide largely without knowledge of the consequences for native ecosystems. Although trout have been introduced widely throughout the Sierra Nevada of California, U.S.A., fishless streams and their communities of native invertebrates persist in some high elevation areas, providing an opportunity to study the effects of trout introductions on natural fishless stream communities. 2. We compared algal biomass and cover, organic matter levels and invertebrate assemblages in 21 natural fishless headwater streams with 21 paired nearby streams containing stocked trout in Yosemite National Park. 3. Although environmental conditions and particulate organic matter levels did not differ between the fishless and trout streams, algal biomass (as chlorophyll a concentration) and macroalgal cover were, on average, approximately two times and five times higher, respectively, in streams containing trout. 4. There were no differences in the overall densities of invertebrates in fishless versus paired trout streams; however, invertebrate richness (after rarefaction), evenness, and Simpson and Shannon diversities were 10–20% higher in fishless than in trout streams. 5. The densities of invertebrates belonging to the scraper‐algivore and predator functional feeding guilds were higher, and those for the collector‐gatherer guild lower, in fishless than trout streams, but there was considerable variation in the effects of trout on specific taxa within functional feeding groups. 6. We found that the densities of 10 of 50 common native invertebrate taxa (found in more than half of the stream pairs) were reduced in trout compared to fishless streams. A similar number of rarer taxa also were absent or less abundant in the presence of trout. Many of the taxa that declined with trout were conspicuous forms (by size and behaviour) whose native habitats are primarily high elevation montane streams above the original range of trout. 7. Only a few taxa increased in the presence of trout, possibly benefiting from reductions in their competitors and predators by trout predation. 8. These field studies provide catchment‐scale evidence showing the selective influence of introduced trout on stream invertebrate and algal communities. Removal of trout from targeted headwater streams may promote the recovery of native taxa, community structure and trophic organisation.     

Ecosystem-level evidence for top-down and bottom-up control of production in a grassland stream system

Ecosystem-wide effects of introduced brown trout (Salmo trutta L.) and native river galaxias (Galaxiaseldoni McDowall) were studied by analysing ecosystem production budgets for two adjacent tributaries of a grassland stream-system in the South Island of New Zealand. One tributary was inhabited by brown trout, the other by river galaxias. No other fish species were present in either stream. The budget for the river galaxias stream indicated little top-down control of invertebrates by fish predation (river galaxias consumed ∼18% of available prey production). A large proportion of annual net primary production was required to support production by invertebrates (invertebrates consumed an average of ∼75% of available primary production), and mean surplus primary production (i.e. not consumed) was not significantly different from zero. Primary and secondary production were presumably mutually limiting in this system (i.e. controlled by simultaneous top-down and bottom-up mechanisms). In contrast, the budget for the brown trout stream indicated extreme top-down control of invertebrate populations by fish predation; essentially all invertebrate production (∼100%) was required to support trout production. Invertebrate production required only a minor portion of annual net primary production (∼21%) and primary production was presumably controlled by mechanisms other than grazing (e.g. sloughing, nutrient limitation). Predatory invertebrates had little quantitative effect on prey populations in either stream. Recent experimental studies of invertebrate behaviour, fish behaviour, and food-web structure in New Zealand streams with physically stable channels indicate that a trophic cascade should be observed in streams inhabited by brown trout, in contrast to those inhabited by native fish. The results reported here provide ecosystem-level evidence supporting this prediction. Received: 10 March 1997 / Accepted: 12 December 1997     

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.     

Diet partitioning in sympatric Atlantic salmon and brown trout in streams with contrasting riparian vegetation

Prey intake by Atlantic salmon Salmo salar and brown trout Salmo trutta was measured across different riparian vegetation types: grassland, open canopy deciduous and closed canopy deciduous, in upland streams in County Mayo, Western Ireland. Fishes were collected by electrofishing while invertebrates were sampled from the benthos using a Surber sampler and drifting invertebrates collected in drift traps. Aquatic invertebrates dominated prey numbers in the diets of 0+ year Atlantic salmon and brown trout and 1+ year Atlantic salmon, whereas terrestrial invertebrates were of greater importance for diets of 1+ and 2+ year brown trout. Terrestrial prey biomass was generally greater than aquatic prey for 1+ and 2+ year brown trout across seasons and riparian types. Prey intake was greatest in spring and summer and least in autumn apart from 2+ year brown trout that sustained feeding into autumn. Total prey numbers captured tended to be greater for all age classes in streams with deciduous riparian canopy. Atlantic salmon consumed more aquatic prey and brown trout more terrestrial prey with an ontogenetic increase in prey species richness and diversity. Atlantic salmon and brown trout diets were most similar in summer. Terrestrial invertebrates provided an important energy subsidy particularly for brown trout. In grassland streams, each fish age class was strongly associated with aquatic, mainly benthic invertebrates. In streams with deciduous riparian canopy cover, diet composition partitioned between conspecifics with older brown trout associated with surface drifting terrestrial invertebrates and older Atlantic salmon associated with aquatic invertebrates with a high drift propensity in the water column and 0+ year fish feeding on benthic aquatic invertebrates. Deciduous riparian canopy cover may therefore facilitate vertical partitioning of feeding position within the water column between sympatric Atlantic salmon and brown trout. Implications for riparian management are discussed.     

Local disturbance history affects patchiness of benthic river algae

1. Recent research has shown that high‐flow events in streams leave a small‐scale mosaic of bed patches that have experienced scouring, sediment deposition (fill), or remained stable. Few studies have investigated if this ‘local disturbance history’ contributes to the patchy distribution of benthic organisms in streams and rivers. 2. In the present research, we demonstrate that local disturbance history in a mid‐sized river can have both short‐ and long‐term effects on epilithic algae. Chains buried vertically in the substratum of the river bed (236 in a 800‐m reach) indicated that two floods (return periods ≤1 year) caused a mosaic of bed patches with different disturbance histories. Once after the first and twice after the second flood, we sampled epilithic algae (mainly diatoms) in replicate patches that had been scoured, filled, or remained stable during the respective event. Algal biomass and cell density per substratum area were determined. 3. Three months after the first flood, algal biomass, total diatom density, diatom taxon richness, and densities of six of nine most common taxa were highest in fill patches. Six days after the second flood, biomass was highest in stable patches, indicating a refugium function of these patches. The refugium patches consisted of average‐sized stones, in contrast to previous studies of flood refugia for benthic algae in which these refugia were always large and/or immobile substrata. Four weeks after the second flood, diatoms tended to be most abundant in scour patches. With one exception, these differences between patch types could not be attributed to differences in local near‐bed current velocity or water depth. 4. The effects of disturbance history were more complex than a simple refugium function of stable patches because algal patterns changed with time since the last disturbance, possibly depending on the successional state of the algal mats.     

The role of trout in stream food webs: integrating evidence from field surveys and experiments

1. We evaluated the effects of brown trout on boreal stream food webs using field surveys and enclosure/exclosure experiments. Experimental results were related to prey preference of uncaged trout in the same stream, as well as to a survey of macroinvertebrate densities in streams with vs. without trout. Finally, we assessed the generality of our findings by examining salmonid predation on three groups of macroinvertebrate prey (chironomid midges, epibenthic grazers, invertebrate predators) in a meta-analysis. 2. In a preliminary experiment, invertebrate predators showed a strong negative response to trout, whereas chironomids benefited from trout presence. In the main experiment, trout impact increased with prey size. Trout had the strongest effect on invertebrate predators and cased caddis larvae, whereas Baetis mayfly and chironomid larvae were unaffected. Trout impact on the largest prey seemed mainly consumptive, because prey emigration rates were low and independent of fish presence. Despite strong effects on macroinvertebrates, trout did not induce a trophic cascade on periphyton. Uncaged trout showed a strong preference for the largest prey items (predatory invertebrates and aerial prey), whereas Baetis mayflies and chironomids were avoided by trout. 3. Densities of invertebrate predators were significantly higher in troutless streams. Baetis mayflies also were less abundant in trout streams, whereas densities of chironomids were positively, although non-significantly, related to trout presence. Meta-analysis showed a strong negative impact of trout on invertebrate predators, a negative but variable impact on mobile grazers (mainly mayfly larvae) and a slightly positive impact on chironomid larvae. 4. Being size-selective predators, salmonid fishes have a strong impact on the largest prey types available, and this effect spans several domains of scale. Discrepancies between our experimental findings and those from the field survey and meta-analysis show, however, that for most lotic prey, small-scale experiments do not reflect fish impact reliably at stream-wide scales. 5. Our findings suggest that small-scale experiments will be useful only if the experimental results are evaluated carefully against natural history information about the experimental system and interacting species across a wide array of spatial scales.     

Interactions between fish,grazing invertebrates and algae in a New Zealand stream: a trophic cascade mediated by fish-induced changes to grazer behaviour?

Experiments in laboratory stream channels compared the behaviour of Deleatidium mayfly nymphs in the absence of fish with that in the presence of either native common river galaxias (Galaxias vulgaris Stokell) or introduced brown trout (Salmo trutta L.). Galaxias present similar predation risks to prey during day and night but are more active at night. Whereas, trout present a higher predation risk during the day. Deleatidium maintained a fixed nocturnal drift periodicity that is characteristic of streams containing visually feeding fish regardless of the nature of the predation regime presented in the laboratory. However, the number on the substratum surface, and therefore able to graze algae, was lower when fish were present than when they were absent. The number was lower during the day in the presence of trout, when they present the highest predation risk, and lower during the night compared to the day in trials with galaxias when galaxias activity disturbs Deleatidium from the substratum. Increases in the probability of Deleatidium leaving a patch, reductions in the proportion of mayflies on high quality patches and reductions in the distance travelled from refuge also reflected variations in the predation regime. Similar differences in positioning were observed under the same predation regimes in in situ channels in the Shag River and these were associated with differences in algal biomass. Algal ash-free dry mass (AFDM) and chlorophyll a (chl a) were higher on the tops of cobbles when fish were present. Fish also affected the biomass and the distribution of algae on cobbles as AFDM and chl a were higher on the sides of cobbles from channels with trout compared to those with galaxias. Changes in grazing behaviour, caused by predator avoidance, are likely to have been responsible for differences in algal biomass because no significant differences were detected between treatments in the biomass of Deleatidium or of total invertebrates.     

Scale-dependant control of motile epifaunal community structure along a coral reef fishing gradient

Large-scale fishing is mostly conducted using towed gears that reduce the biomass and diversity of benthic invertebrates. However, it is impossible to differentiate between the physical disturbance effect of towed gears from the effect of fish predator removal upon benthic invertebrate communities. Here we explore the impact of fish removal alone on the community structure of small motile coral reef invertebrates (epifauna) along a subsistence fishing intensity gradient in the Lau group, Fiji. We deployed settlement plates at three areas in each of six fishing grounds and examined the density and class richness of the motile epifaunal communities and the associated algal communities in relation to the structure of fish and benthic communities. Motile epifaunal density was unrelated to fishing intensity. However, at smaller inter-area scale (0.5-10 km) motile epifaunal density was negatively related to plate algal biomass, whereas at the larger inter-fishing-ground scale (4-180 km) motile epifaunal density was positively related to the rugosity (substrate complexity) of the surrounding benthos. The class richness and diversity (Margalef's d) of motile epifaunal communities were negatively related to fishing intensity, but unrelated to grazing intensity, rugosity or algal biomass at either scale. Benthic community structure varied significantly with fishing intensity; hard-coral cover was lower and turf-algal cover was higher at high fishing pressure. The variation in benthic community structure was associated with variation in fish community structure, which in turn varied with fishing intensity. Motile epifaunal community structure upon plates was linked to the structure of the surrounding benthic community, but was not directly linked to the plate algal community. We suggest the decline in richness of the motile epifauna community along the fishing gradient is attributable to either to exploiter-mediated coexistence or the reduction in ‘habitat quality’ of the surrounding benthos. At the large spatial scale substrate complexity is the key determinant of motile epifaunal density, suggesting predation by fishes plays an important structuring role at this scale. Assuming that rugosity is inversely related to predation risk then this study represents the first evidence for spatial-dependence on the top-down (predation) vs. bottom-up (algal biomass) control of community structure. We argue fisheries exploitation, in the absence of a physical disturbance can negatively influence motile epifaunal community structure at large spatial scales.     

The effect of two scales of habitat architecture on benthic grazing in a river

1. This experiment studied the effects of differing levels of the complexity of substratum architecture at two spatial scales on the distribution and abundance of benthic algae and invertebrates, and the strength of the trophic interaction between invertebrate grazers and algae. Some estimates of the effects on invertebrate colonization rates were also made. 2. Four levels of microhabitat architectural complexity were created using artificial substrata (clay tiles) and placed in Mountain River, Tasmania, in two riffle types (bedrock and boulder-cobble) of differing large-scale substratum complexity. After a colonization period, invertebrate grazers were removed from half the tiles to measure the effects of grazing. Invertebrates on the tiles were also counted and identified. At the end of the experiment, algae were removed from the tiles and analysed for chlorophyll a. 3. Invertebrate grazers did not reduce algal biomass during the experiment, and microhabitat-scale architecture influenced algal biomass more strongly than riffle-scale architecture. Highly complex microhabitat architecture increased algal biomass by providing more surface area, but once standardized for surface area, algal biomass decreased as the complexity of microhabitat architecture increased. 4. Microhabitat-scale architecture was also predominant in determining invertebrate density and the identity of the dominant grazer species. In contrast to algal biomass, invertebrate densities and species density increased with the complexity of microhabitat architecture, suggesting that refuges from flow (and possibly predation) were as important to river invertebrates as the distribution of their food source. 5. Riffle-scale architecture had some effect on the colonization of two slow-moving grazer taxa, but, overall, the colonization processes of slow-moving grazers were determined mostly by the complexity of microhabitat-scale architecture.     

Local disturbance history and habitat parameters influence the microdistribution of stream invertebrates

1. We investigated the effects of local disturbance history and habitat parameters (abiotic and biotic) on the microdistribution of benthic invertebrates during several floods in two streams, the Schmiedlaine in Germany (four events) and the Kye Burn in New Zealand (two events). 2. Bed movement patterns were quantified using metal‐link scour chains. Before and after each flood, quantitative invertebrate samples were taken from replicate bed patches that had experienced sediment scour, fill or remained stable. 3. Patterns of invertebrate density in the different bed stability types (i.e. scour, fill, stable) varied between floods, sampling dates and streams, but invertebrate density was highest in stable patches in >50% of all the patch type effects detected and lowest in fill patches in 75% of all detected effects. Stable bed patches acted as a refugium for Liponeura spp. and Leuctra spp. in the Schmiedlaine and for Hydracarina and Deleatidium spp. in the Kye Burn. 4. Averaged across both streams, only near‐bed current velocity was correlated with invertebrate distribution on the streambed more often than disturbance history. In the Kye Burn, disturbance history and water depth were the most influential habitat parameters. 5. Our results suggest that a thorough understanding of the microdistribution of benthic invertebrates requires knowledge of disturbance history, as well as more readily measured habitat parameters such as current velocity or water depth.     

Influence of fishes on macroinvertebrate communities and insect emergence production in intermittent stream refuges

1. Drying intermittent stream networks often have permanent water refuges that are important for recolonisation. These habitats may be hotspots for interactions between fishes and invertebrates as they become isolated, but densities and diversity of fishes in these refuges can be highly variable across time and space.
2. Insect emergence from streams provides energy and nutrient subsidies to riparian habitats. The magnitude of such subsidies may be influenced by in-stream predators such as fishes.
3. We examined whether benthic macroinvertebrate communities, emerging adult insects, and algal biomass in permanent grassland stream pools differed among sites with naturally varying densities of fishes. We also manipulated fish densities in a mesocosm experiment to address how fishes might affect colonisation during recovery from hydrologic disturbance.
4. Fish biomass had a negative impact on invertebrate abundance, but not biomass or taxa richness, in natural pools. Total fish biomass was not correlated with total insect emergence in natural pools, but orangethroat darter (Etheostoma spectabile) biomass was inversely correlated with emerging Chironomidae biomass and individual midge body size. The interaction in our models between predatory fish biomass and date suggested that fishes may also delay insect emergence from natural pools, altering the timing of aquatic–terrestrial subsidies.
5. There was an increase over time in algal biomass (chlorophyll-a) in mesocosms, but this did not differ among fish density treatments. Regardless, fish presence in mesocosms reduced the abundance of colonising insects and total invertebrate biomass. Mesocosm invertebrate communities in treatments without fishes were characterised by more Chironomidae, Culicidae, and Corduliidae.
6. Results suggest that fishes influence invertebrates in habitats that represent important refuges during hydrologic disturbance, hot spots for subsidy exports to riparian food webs, and source areas for colonists during recovery from hydrologic disturbance. Fish effects in these systems include decreasing invertebrate abundance, shifting community structure, and altering patterns of invertebrate emergence and colonisation.

     

Microform bed clusters: are they preferred habitats for invertebrates in a flood‐prone stream?

SUMMARY 1. Microform bed clusters are bedform microunits in streams. They consist of an obstacle clast (a large surface stone) against which other stones are stacked in a specific manner. Stream ecologists have suggested that these clusters are better flood refugia and more valuable habitats for invertebrates than single surface stones because of their higher stability during high‐flow events and their greater diversity of microhabitats available for colonisation. 2. To test these predictions in the Schmiedlaine, a flood‐prone prealpine stream, we sampled invertebrates on clustered and single stones on a total of seven occasions including before and after a moderate spate, two major floods and a minor spate. We also monitored surface particle stability during the second major flood and the minor spate. 3. Before the second flood (return period 1–2 years), we determined the exact positions of 60 clusters (60 obstacle clasts with 136 associated smaller particles) and 50 single stones in a 250‐m reach. We sampled the fauna on stones in the clusters, on well‐embedded single stones, and on loose single stones. Only six obstacle clasts and one single stone remained after the flood, implying movement of almost the entire surface layer. Therefore, we sampled the invertebrates on stones in newly formed clusters, on well‐embedded and single loose stones 3 and 11 days after the flood. The minor spate, in contrast to the second flood, moved only one of 495 monitored surface particles. 4. Three days after the flood, invertebrate density on clustered stones and both single stone categories were similar and equalled 34% of the mean pre‐flood density. Eight days later, density had almost doubled. The relatively high survival and rapid recovery suggest that invertebrates found refugia during the flood. However, the very low stability of clusters and single stones implied that surface particles were unimportant as refugia. 5. Total invertebrate density and taxon richness were never higher on clustered than on single stones (regardless of the timing of sampling relative to the last previous high‐flow event). Densities of the seven most common taxa and invertebrate community structure were also generally similar between particle types. We conclude that microform bed clusters cannot be regarded as more valuable invertebrate habitats than single surface stones in the Schmiedlaine.     

Plant—herbivore interactions in streams near Mount St Helens

1. In four separate field experiments near Mount St Helens (Washington, U.S.A.) during 1986, the grazing effects of two large benthic herbivores, tadpoles of the tailed frog Ascaphus truei and larvae of the caddisfly Dicosmoecus gilvipes, were investigated using streamside channels and in-stream manipulations. In the experimental channels, abundances of periphyton and small benthic invertebrates declined significantly with increasing density of these larger herbivores. 2. In eleven small, high-gradient streams affected to varying degrees by the May 1980 eruption, in-stream platforms were used to reduce grazing by A, truei tadpoles on tile substrates. Single platforms erected in each tributary and compared to grazed controls revealed only minor grazing effects, and no significant differences among streams varying in disturbance intensity (and, consequently, tadpole density). However, results probably were confounded by high variability among streams in factors other than tadpole abundance. 3. Grazing effects were further examined in two unshaded streams with different tadpole densities, using five platforms per stream. In the stream with five tadpoles m^?2, grazing reduced periphyton biomass by 98% and chlorophyll a by 82%. In the stream lacking tadpoles, no significant grazing effects were revealed. Low algal abundance on both platforms and controls, and high invertebrate density in that stream (c. 30000m^?2) suggests that grazing by small, vagile invertebrates was approximately equivalent to that of tadpoles. 4. The influence of large benthic herbivores on algal and invertebrate communities in streams of Mount St Helens can be important, but reponses vary spatially in relation to stream disturbance history, local environmental factors, and herbivore distributional patterns and abundance.     

The influence of moss on grazers in high‐altitude streams: food,refuge or both?

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Brown trout and food web interactions in a Minnesota stream

1. We examined indirect, community‐level interactions in a stream that contained non‐native brown trout (Salmo trutta Linnaeus), native brook trout (Salvelinus fontinalis Mitchill) and native slimy sculpin (Cottus cognatus Richardson). Our objectives were to examine benthic invertebrate composition and prey selection of fishes (measured by total invertebrate dry mass, dry mass of individual invertebrate taxa and relative proportion of invertebrate taxa in the benthos and diet) among treatments (no fish, juvenile brook trout alone, juvenile brown trout alone, sculpin with brook trout and sculpin with brown trout). 2. We assigned treatments to 1 m² enclosures/exclosures placed in riffles in Valley Creek, Minnesota, and conducted six experimental trials. We used three designs of fish densities (addition of trout to a constant number of sculpin with unequal numbers of trout and sculpin; addition of trout to a constant number of sculpin with equal numbers of trout and sculpin; and replacement of half the sculpin with an equal number of trout) to investigate the relative strength of interspecific versus intraspecific interactions. 3. Presence of fish (all three species, alone or in combined‐species treatments) was not associated with changes in total dry mass of benthic invertebrates or shifts in relative abundance of benthic invertebrate taxa, regardless of fish density design. 4. Brook trout and sculpin diets did not change when each species was alone compared with treatments of both species together. Likewise, we did not find evidence for shifts in brown trout or sculpin diets when each species was alone or together. 5. We suggest that native brook trout and non‐native brown trout fill similar niches in Valley Creek. We did not find evidence that either species had an effect on stream communities, potentially due to high invertebrate productivity in Valley Creek.     

What predicts the use by brook trout (Salvelinus fontinalis) of terrestrial invertebrate subsidies in headwater streams?

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Effects of trout on the diel periodicity of drifting in baetid mayflies

Some benthic invertebrates in streams make frequent, short journeys downstream in the water column (=drifting). In most streams there are larger numbers of invertebrates in the drift at night than during the day. We tested the hypothesis that nocturnal drifting is a response to avoid predation from fish that feed in the water column during the day. We surveyed diel patterns of drifting by nymphs of the mayfly Baetis coelestis in several streams containing (n=5) and lacking (n=7) populations of rainbow trout, Oncorhynchus mykiss. Drifting was more nocturnal in the presence of trout (85% of daily drift occurred at night) than in their absence (50% of daily drift occurred at night). This shift in periodicity is due to reduced daytime drifting in streams with trout, because at a given nighttime drift density, the daytime drift density of B. coelestis was lower in streams occupied by trout than in troutless streams. Large size classes of B. coelestis were underrepresented in the daytime drift in trout streams compared to nighttime drift in trout streams, and to both day and night drift in troutless streams. Differences in daytime drift density between streams with and without trout were the result of differences in mayfly drift behaviour among streams because predation rates by trout were too low to significantly reduce densities of drifting B. coelestis. We tested for rapid (over 3 days) phenotypic responses to trout presence by adding trout in cages to three of the troutless streams. Nighttime drifting was unaffected by the addition of trout, but daytime drift densities were reduced by 28% below cages containing trout relative to control cages (lacking trout) placed upstream. Drift responses were measured 15 m downstream of the cages suggesting that mayflies detected trout using chemical cues. Overall, these data support the hypothesis that infrequent daytime drifting is an avoidance response to fish that feed in the water column during the day. Avoidance is more pronounced in large individuals and is, at least partially, a phenotypic response mediated by chemical cues.     

Habitat structure,resources and diversity: the separate effects of surface roughness and macroalgae on stream invertebrates

Habitat structure has pervasive effects on community composition and diversity, with physically complex habitats often containing more species than physically simple ones. What factors or mechanism drive this pattern is little understood, but a complicating problem is that different sources of habitat structure can be confounded in both surveys and experiments. In this study, we carried out an experiment in which two sources of habitat structure, attached macroalgae and substrate surface texture, were separately manipulated to discern their joint and separate effects upon the diversity and composition of colonizing macroinvertebrates in a stony, upland stream. Because stream algae vary markedly in abundance in both space and time, we also sampled the epilithon of stream stones at two spatial scales on eight dates over 2 years to gain some preliminary data on how stream algae vary between individual substrata over time. Experimental substrata had either a smooth (siltstones, sandstones, crystal-poor felsic volcanics, plain paving bricks) or rough (granodiorites, crystal-rich felsic volcanics, sand-blasted paving bricks) surface. We allowed these substrata to be colonized naturally by macroalgae, mostly the filamentous red alga Audouinella hermannii. Half of each of the rough and smooth substrata were selected at random and the macroalgae gently sheared off. All substrata were defaunated with a household insecticide with little field persistence, set out randomly through the study riffle, and invertebrates allowed to colonize them for 14 days. Some substrata were sampled immediately to check the efficacy of faunal and algal removals, which proved to be successful. Experimental results showed that both surface texture and macroalgae increase species richness independently of each other. Surface texture had no effect on densities, while macroalgae increased colonization densities, but rarefaction showed that both sources of habitat structure increased species richness above values expected simply on the basis of the numbers of colonists. However, reference stones with high macroalgal cover had the same species richness as those with low cover, suggesting that the effects of macroalgae on species richness are transient relative to those associated with surface texture. Epilithon samples taken at different times suggest that the magnitude of spatial variation in plant growth alters with time. If plants generally recolonize rough surfaces more quickly than smooth, then the effects of habitat structure on macroinvertebrates ought to be strongest after major disturbances during growing seasons of plants. Received: 1 September 1999 / Accepted: 10 January 2000