Food and space revisited: The role of drift-feeding theory in predicting the distribution,growth, and abundance of stream salmonids

In this paper we review drift-feeding models for stream salmonids. We assess their historical development and current state, and we propose areas for future research. Drift-feeding models serve as the critical input for energetics-based habitat selection and habitat quality models, which have recently begun to see widespread use for predicting salmonid distribution, growth and abundance. We use a bibliometric approach to find drift-feeding model publications, especially those citing three landmark papers that began the quantification of drift feeding by stream fish (Fausch 1984; Hughes and Dill 1990; Hill and Grossman 1993). Subsequent drift-feeding models have largely been built upon these models. Research effort has focused on model development and applications but model testing has been neglected. To date, the only rigorous test of a drift-feeding model (Hughes et al. 2003) identified several limitations and violations of model assumptions. The most important limitation was that prey capture- and gross energy intake rates were overestimated by a factor of two, due largely to poor predictions of prey detection probabilities. Consequences of error in drift-feeding models, and consequently in the habitat selection/quality models that employ them, are greater for applications aimed at predicting growth and abundance than they are for predicting distribution. Research effort on a broad front is needed to advance both drift-feeding models and habitat selection/quality models, including: further development of drift-foraging theory, revision and testing of drift-feeding models (specifically new, functional prey detection and interception sub-models), and revision of habitat selection/quality models to incorporate spatial, temporal, and flow-dependent variation in drift concentration.     

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     

Temporal consistency in the long-term spatial distribution of macroinvertebrate drift along a stream reach

Previous studies of the spatial pattern of stream invertebrate drift have focused on spatial variation at microhabitat scales or landscape scales, or temporal variation over diel or seasonal scales. None have examined consistency in spatial variation over longer time scales (>1 year). This study examined invertebrate drift density and composition at fixed locations (terminal ends of 10 riffles) each month at day and night along a 1 km reach of a 2nd order stream over a period of nearly 2 years. Consistent differences in the density of macroinvertebrate drift between riffles over 2 years were observed. The only habitat characteristic observed to be related to invertebrate drift density was the length and size of riffles above sampling sites, with larger and longer riffles producing the highest drift densities. Consistent differences in the supply of drifting macroinvertebrates along a stream reach may have implications for the supply of colonists to substrate patches and the profitability of feeding positions for drift-feeding fish and other predators. Handling editor: D. Dudgeon     

Ontogenetic shift in nocturnal microhabitat selection by giant kokopu in a New Zealand stream

Nocturnal microhabitat selection in relation to size was examined for a New Zealand drift-feeding freshwater fish, the giant kokopu Galaxias argenteus in three coastal Otago streams. Evidence for ontogenetic shifts in microhabitat selection were observed, particularly with respect to water velocity and depth. Small (<80 mm) giant kokopu tended to occur in shallow backwaters adjacent to fast flowing water. In contrast, larger individuals (≥80 mm) were most commonly observed in deeper, slower flowing pools. These patterns of habitat selection are most likely associated with the trade-offs involved with the selection of optimal habitats for drift feeding combined with avoidance of intra-specific competition and the threat of predation from larger conspecifies.     

Hydro-morphologically related variance in benthic drift and its importance for numerical habitat modelling

Numerical fish-habitat modelling on various scales is considered to be state of the art in river management. However, most of the concepts applied use steady-state hydraulic parameters such as flow velocity and water depth. Herein we present analysis and discussion of the possibility of including a drift-feeding parameter (SI_F) into habitat evaluations based on multiplying suitability indices. “Sources” and “sinks” of benthic drift were identified according to both the zero-crossing and hydraulic-threshold methods in an alpine gravel-bed river. Minor differences could be determined between the two methods in a well-developed riffle–pool section. Macroinvertebrates, used for simulating benthic drift, were collected by multi-habitat sampling and appraised according to their critical threshold (τ _cr) for motion on the bed surface and sinking velocity (v _s). The findings of the calculation of drift rates using one- (1D) and two-dimensional (2D) hydrodynamic numerical models highlight a specification of best feeding position for drift-feeding fish (i.e. brown trout, grayling) considering the SI_F parameter. Riffle–pool sequences are characteristic of pristine alpine streams; our findings underline their importance as production (riffles) and consumption areas (pools) in terms of holistic river function. Moreover, the results indicate that (artificial) lateral obstruction (e.g. dams) may lead to a reduced transport rate of benthic organisms due to low bottom shear stress (<0.25 N m⁻²). Thus, deposition of drifting macroinvertebrates occurs in backwaters, with downstream impacts on benthic and fish communities.     

Woody debris and terrestrial invertebrates – effects on prey resources for brown trout (Salmo trutta) in a boreal stream

Intensive forestry and other activities that alter riparian vegetation may disrupt the connectivity and the flux of energy between terrestrial and aquatic habitats and have large effects on biota, especially in small streams. We manipulated the amount of in-stream wood and the flux of terrestrial invertebrate subsidies to determine how these factors affected potential food resources for drift-feeding brown trout (Salmo trutta ) in a boreal Swedish forest stream. Specifically, we followed the effects on the abundance of aquatic and terrestrial invertebrate fauna from June to August 2007. The treatments were 1) addition of wood, unmanipulated terrestrial invertebrate inputs, 2) reduction of terrestrial invertebrate inputs (using canopy covers), no addition of wood, 3) unmanipulated ambient conditions, 4) simultaneous addition of wood and reduction of terrestrial invertebrate inputs. Added wood resulted in greater biomass of aquatic invertebrate biomass, and both input and drift of terrestrial invertebrates were reduced by canopy covers. In terms of total potential prey biomass, the addition of wood with ambient levels of terrestrial invertebrate inputs had the highest standing crop of benthic, wood-living and terrestrial invertebrates combined, whereas the treatment with reduced terrestrial input and no wood added had the lowest standing crop. Our study indicates that forest practices that both reduce the recruitment of wood and the input of terrestrial invertebrates to small streams have negative effects on prey availability for drift-feeding brown trout. The positive effects of wood addition on biomass of aquatic macroinvertebrates may partly compensate for the negative effects of reduced terrestrial invertebrate subsidies.     

Not all drift feeders are trout: a short review of fitness-based habitat selection models for fishes

Currently, there are few mechanistic fitness-based habitat selection models for stream fishes and most models used by management agencies focus on physical habitat alone. In this review, I describe the historical development and the status of mechanistic, fitness-based, habitat selection models for both water column (i.e., drift-feeding) and benthic stream fishes focusing on North America. Although the majority of drift feeders are not salmonids, most mechanistic habitat selection models have been developed and tested only in this group of fishes, likely due to their substantial economic importance. I review the fitness-based microhabitat selection model of Grossman et al. (Ecol Fresh Fish 11:2–10, 2002), which has been tested in both a salmonid and multiple cyprinid species. The model predicts optimal focal point velocities for drift feeders based on prey capture success–velocity relationships and does not include physiological costs, which are logistically difficult to quantify. In addition, I discuss mechanistic, fitness-based models used to predict microhabitat (i.e., patch) selection in benthic fishes. For both basic scientific and management/conservation perspectives, it is important to quantify habitat choice in fishes using mechanistic, fitness-based criteria.     

Invertebrate responses to short-term water abstraction in small New Zealand streams

1. Small permanent streams are coming under increasing pressure for water abstraction. Although these abstractions might only be required on a short‐term basis (e.g. summer time irrigation), the highest demand for water often coincides with seasonal low flows. 2. We constructed weirs and diversions that reduced discharge in three small streams (<4 m width) to test the hypotheses that short‐term water abstractions would decrease habitat availability and suitability for invertebrates, resulting in increased invertebrate drift, reduced taxonomic richness and decreased benthic invertebrate densities. 3. We sampled benthic invertebrates, invertebrate drift and periphyton at control (upstream) and impact (downstream) sites on each stream before and during 1 month of discharge reduction. 4. Discharge decreased by an average of 89–98% at impact sites and wetted width decreased by 24–30%. Water depth decreased by 28–64% while velocity decreased by 50–62%. Water conductivity, temperature and dissolved oxygen showed varying responses to flow reduction among the three streams, whereas algal biomass and pH were unaffected in all streams. 5. The densities of invertebrate taxa tended to increase in the impact reaches of these streams, even though invertebrate drift increased at impact sites in the first few days following discharge reduction. There were a higher proportion of mayflies, stoneflies and caddisflies at the impact site on one stream after flow reduction. There were no changes to the number of taxa or species evenness at impact sites. 6. Our results suggest that for these small streams, the response of invertebrates to short‐term discharge reduction was to accumulate in the decreased available area, increasing local invertebrate density.     

The relation between invertebrate drift and two primary controls,discharge and benthic densities,in a large regulated river

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Nocturnal drift of mayfly nymphs as a post-contact antipredator mechanism

1. The predominantly nocturnal constrained drift of stream invertebrates is commonly regarded as a behaviour that avoids encounters with visually foraging fish in the water column. The alternative explanation, that drift peaks are caused by bottom-feeding, nocturnal predators, has rarely been tested.
2. We examined these hypotheses by collecting invertebrate drift in five streams in northern Finland: one with brown trout ( Salmo trutta , a drift-feeding fish), one with alpine bullhead ( Cottus poecilopus , a benthic fish), one with both species, and two fishless streams.
3. Drift by Baetis mayflies was aperiodic or slightly diurnal in both fishless streams on all sampling occasions. In contrast, drift was nocturnal in streams with trout and, to a lesser extent, in the stream with bullhead. Non-dipteran prey drifted mainly nocturnally in all streams with fish, whereas Diptera larvae were less responsive to the presence of fish.
4. In laboratory experiments, bullheads were night-active, causing a much higher frequency of drift by touching Baetis at night than during the day. Thus, increased nocturnal drift may serve to avoid both visual predators (a pre-contact response) and benthic fish (a post-contact response). In streams with bottom-feeding fish, nocturnal drift should be caused by increased drift by night rather than by reduced drift by day.     

Comparing correlative and bioenergetics‐based habitat suitability models for drift‐feeding fishes

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Day and night drift-feeding by juvenile salmonids at low water temperatures

Drift-feeding salmonids in boreal streams face temperatures below physical optima for extensive periods of the year. Because juvenile salmonids react to low water temperatures by becoming nocturnal, knowledge about their foraging ability at low light intensities in cold water is needed to accurately estimate energy intake during non-summer conditions. In a laboratory stream channel, we studied temperature effects on the drift-feeding behaviour of juvenile Atlantic salmon, brown trout, and European grayling in simulated daylight and moonlight at temperatures ranging from 2 °C to 11 °C. Prey capture probability was positively related to temperature, but the temperature dependence did not agree with predictions of the Metabolic Theory of Ecology. Furthermore, reaction distance was positively related to temperature for the three species, which may be one of the underlying mechanisms responsible for the temperature effects on prey capture probability. Overall, the three species had similar capture rates at the different temperature and light levels, although there were species differences. European grayling had a slightly higher prey capture probability than brown trout, and brown trout had a shorter reaction distance than Atlantic salmon and European grayling. These results have implications for both energetics-based drift-foraging theory and for studies of winter ecology.     

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.     

Simple predictions of instream habitat model outputs for target fish populations

1. Bottom-up approaches based on individual behaviour can help to identify key variables influencing populations at larger scales. Instream habitat models have been developed to predict the consequences, for populations in stream reaches, of fish preferences for particular hydraulic conditions observed at the scale of individuals. Conventional instream habitat models (e.g. PHABSIM) predict habitat values for species or life stages in reaches, and their changes with discharge. Despite their worldwide use, they have been subject to continuing criticism and have been mainly limited to site-specific case studies.
2. We ran conventional instream habitat models in 58 French stream reaches dominated by brown trout. Using non-linear mixed effect models, we demonstrated that the outputs of instream habitat models (habitat values for three trout life stages and five other species) are predictable from average characteristics of reaches (discharge, depth, width and bed particle size).
3. Our models closely reflect variations in habitat values within-reaches (with discharge) and between-reaches. Within-reach changes are linked to the Reynolds number of reaches, while between-reach changes depend mainly on the Froude number at median daily discharge. These two dimensionless variables combine discharge, mean depth and mean width of reaches. Independent model validations showed robust model predictions that are consistent with studies of habitat values for brown trout made in larger streams from western North America.
4. Our results contribute to identifying the main hydraulic variables governing estimates of fish habitat values. They should facilitate habitat studies in multiple streams, at the basin or larger scales, while reducing their cost. They should enhance the biological validation of habitat model predictions, which remains critical.     

Species traits and channel architecture mediate flow disturbance impacts on invertebrate drift

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The dynamics of terrestrial invertebrate inputs to the food web of a small salmon river

The qualitative composition and quantitative characteristics of the syrton allochthonous fraction in one of the rivers of the Russian Far East are examined. It is assumed that the daily dynamics of terrestrial invertebrate drift is determined by the specificity of their daily migrations on land. A counterargument to the theory of the zoobenthos nighttime drift strategy as a way to avoid drift-feeding predators is suggested.     

Mechanisms of drift-feeding behavior in juvenile Chinook salmon and the role of inedible debris in a clear-water Alaskan stream

Drift-feeding fish are challenged to discriminate between prey and similar-sized particles of debris, which are ubiquitous even in clear-water streams. Spending time and energy pursuing debris mistaken as prey could affect fish growth and the fitness potential of different foraging strategies. Our goal was to determine the extent to which debris influences drift-feeding fish in clear water under low-flow conditions when the distracting effect of debris should be at a minimum. We used high-definition video to measure the reactions of drift-feeding juvenile Chinook salmon (Oncorhynchus tshawytscha) to natural debris and prey in situ in the Chena River, Alaska. Among all potential food items fish pursued, 52 % were captured and quickly expelled from the mouth, 39 % were visually inspected but not captured, and only 9 % were ingested. Foraging attempt rate was only moderately correlated with ingestion rate (Kendall’s τ?=?0.55), raising concerns about the common use of foraging attempts as a presumed index of foraging success. The total time fish spent handling debris increased linearly with foraging attempt rate and ranged between 4 and 25 % of total foraging time among observed groups. Our results help motivate a revised theoretical view of drift feeding that emphasizes prey detection and discrimination, incorporating ideas from signal detection theory and the study of visual attention in cognitive ecology. We discuss how these ideas could lead to better explanations and predictions of the spatial behavior, prey selection, and energy intake of drift-feeding fish.     

Seasonal and diel patterns of invertebrate drift in different alpine stream types

1. We examined the seasonal and diel patterns of invertebrate drift in relation to seston and various habitat characteristics in two each of four different kinds of alpine streams [rhithral (snow‐fed) lake outlets, rhithral streams, kryal (glacial‐fed) lake outlets and kryal streams]. Samples were collected at four times of the day (dawn, midday, dusk and midnight) during three seasons (spring, summer and autumn). 2. Habitat characteristics differed mainly between rhithral and kryal sites, with the latter having higher discharge and turbidity, lower water temperature, and higher concentrations of ammonium, and particulate and soluble reactive phosphorus. Seasonality in habitat characteristics was most pronounced for kryal streams with autumn samples being more similar to rhithral sites. 3. The concentration of seston was lowest in the glacial‐influenced lake outlets and slightly higher in the stream sites; no seasonal or diel patterns were evident. 4. The density of drifting invertebrates averaged less than 100 m^?3 and was lowest (<10 m^?3) at three of the four kryal sites. Taxon richness and diversity were lowest at rhithral lake outlets. Chironomidae dominated the drift as well as benthic communities and <30% of benthic taxa identified were found in the drift. 5. Drifting invertebrates showed no consistent seasonal pattern. However, density tended to be highest in spring at rhithral sites and in autumn at kryal sites. No diel periodicity in drift density was found at any site and the lack of diel pattern may be a general feature of high altitude streams. 6. Glacially influenced habitat parameters were a major factor affecting drift in these alpine streams, whereas no clear differences were observed between streams and lake outlets. Our findings indicate that invertebrate drift in alpine streams is primarily influenced by abiotic factors, and therefore, substantially differs from patterns observed at lower altitude.     

Invertebrate mesopredators are larger in streams with fish

Species with complex life cycles (e.g., aquatic larvae, terrestrial adults) are expected to shorten the time spent in the larval stage if mortality risks are high, a trade-off that lowers predation risk at the cost of reduced time for growth and thus smaller adult size. We tested these predictions by comparing the timing of and size at emergence for two relatively large and common invertebrate mesopredator species (Isoperla montana and Rhyacophila vibox) that inhabit small coastal streams, with and without predatory fish, in eastern Canada. Contrary to expectations based on predation risk–foraging trade-off theory, individuals of both invertebrate species tended to be larger rather than smaller in streams with fish than in fishless streams. The patterns were consistent, however, with the expected ecological effects of top predators on food webs, where fish lower abundances of invertebrate mesopredators, increasing resource availability and thus growth rates for the remaining individuals. We conclude that variation among streams in size at emergence is better explained by the impact of fish on resource availability than to behavioural or life history trade-offs occurring under risk of predation.     

A historical perspective on drift foraging models for stream salmonids

Thirty years ago, Fausch (Can J Zool 62:441–451, 1984) proposed a simple model of optimal positions for drift-feeding salmonids in streams, whereby fish maximize their net energy intake (NEI) by selecting focal points in low water velocity near faster currents that deliver abundant drifting invertebrates. The theory was based on earlier observations in artificial and natural streams describing characteristics of salmonid positions and a conceptual model by Chapman (Am Nat 100:345–357, 1966). A test of this simple drift foraging model in a laboratory stream showed that the growth rate of juvenile trout and salmon increased with NEI, and that the rank of NEI at positions held by coho salmon (Oncorhynchus kisutch) correlated nearly perfectly with their rank in the dominance hierarchy. Fausch (1984) inferred from these findings that positions that optimize NEI, within the constraints of the dominance hierarchy, are the resource for which these stream salmonids compete. In turn, the model was used to test the effects of interspecific competition by coho salmon on the foraging positions held by brook trout (Salvelinus fontinalis) and brown trout (Salmo trutta), and these results were used to infer potential effects of the introduced salmon on resident trout in Lake Michigan tributaries. Though the goals for this model were originally modest, it was tested in the field and further refined by Hughes and Dill (Can J Fish Aquat Sci 47:2039–2048, 1990) and others. During the last 20 years, the general theory has been incorporated into other models, which have been applied widely to analyze salmonid distribution and abundance in streams and rivers and used for management and restoration of habitat and flow regimes to benefit these fishes.