Stream drift as a consequence of disturbance by invertebrate predators

Summary We carried out an experimental field study in a Swedish stream in order to determine whether mobile predators enhance the drift of stream insects. We increased the density of nymphs of the predaceous perlid stonefly, Dinocras cephalotes, in an experimental section of a stream up to densities in another more densely populated part of the same stream. The drift of several benthic species increased significantly compared to a control section where D. cephalotes were rare. Experiments carried out in September showed a strongly elevated drift response in nymphs of the mayfly Baetis rhodani only, whereas May experiments resulted in increased drift in B. rhodani as well as the amphipod Gammarus pulex, the stonefly Leuctra fusca, chironomids, and the total number of drifting animals. In September, we found that the drift response of Baetis rhodani to predator disturbance was dependent on the size of mayfly nymphs; small nymphs appeared in greater numbers in the drift nets than did large nymphs. A subsequent laboratory analysis of drift lengths of B. rhodani nymphs supported the hypothesis that small nymphs travel in the drift for longer than do large nymphs, particularly in darkness. We suggest that morphological constraints in vision or swimming performance, or both, cause small nymphs to drift longer. In May, size-dependent drift was less obvious, probably because the size of the nymphs was considerably greater than in September.     

Diel vertical movements by lotic mayfly nymphs under variable predation risk

1. The diel foraging periodicities of two grazing mayfly (Ephemeroptera) nymphs, Heptagenia dalecarlica and Baetis rhodani, under variable fish (European minnow) predation risk were examined in a series of laboratory experiments. 2. Heptagenia dalecarlica were almost exclusively nocturnal in their use of feeding areas on stone tops. There was a sharp increase in the proportion of nymphs out of refuge at nightfall, both in the control and fish-odour treatments. In a treatment with freely moving fish, H. dalecarlica decreased their activity during both the day and night. In feeding trials with three freely foraging minnows, nymphs were completely safe when they had access to refuges beneath stones, whereas almost half the nymphs were consumed when no refuges were available. 3. Baetis rhodani nymphs reduced their use of stone tops when exposed to four caged minnows or a freely moving fish, but this occurred both day and night. In feeding trials, B. rhodani were captured only while in the water column, and their mortality risk was independent of refuge availability. 4. It is suggested that because H. dalecarlica lack efficient post-encounter defences, they must rely on pre-encounter mechanisms to reduce the threat of fish predation. It appears that in order to dwell sympatrically with fish, H. dalecarlica have evolved a coexistence by hiding strategy. In contrast, Baetis are vulnerable to fish attacks only if they enter drift in daylight, and are thus able to remain on stone tops both during the day and at night.     

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

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Long‐term monitoring of the life history of a riverine mayfly Ephemera orientalis (Ephemeroptera: Ephemeridae) in the Han River in Seoul,Korea

Climatic change often affects life history aspects of aquatic insects. Long‐term monitoring was conducted to understand the life history variability of the riverine mayfly Ephemera orientalis in the Han river, where habitat is largely disturbed by summer floods. Water level of the study site, Godeok‐dong area in Seoul, is regulated by the Paldang Dam located approximately 14 km above the study site. E. orientalis often emerged in large numbers around the study area. Larvae were sampled monthly from April 2006 to July 2010 using a Surber sampler (50 × 50 cm, mesh 0.25 mm, two replicates). Additional qualitative sampling was conducted using a hand net. As a result, based on the larval body size distribution, E. orientalis had a univoltine life cycle with two distinct slow growing cohort groups: the S1group emerged during May–June and the S2 group emerged during August–September. Our previous study conducted in Gapyeong stream in Gyeonggi‐do showed that E. orientalis normally had three cohort groups, S‐1 and S‐2 groups and the F‐group (a fast growing cohort developed during May–August). However, the Han River population of E. orientalis lacked such a fast growing cohort. It is most probable that habitat disturbances caused by high water level during the rainy season negatively affected the development of the fast growing cohort (F‐group) in the Han River area. The relationships between monthly E. orientalis population data and water level in the Han River are presented.     

The relative importance of refugia in determining the drift and habitat selection of predaceous stoneflies in a sandy-bottomed stream

Patch structure in sandy, compared to rocky streams, is characterized by isolated snags that can only be colonized by drifting. By measuring drift from patches (snags) of various quality we determined the factors that influence habitat selection and drift of the predaceous stoneflies Acroneuria abnormis and Paragnetina fumosa. The presence of refugia (loose bark and leaf packs) was more important than hunger level and modified the effects of increased predator densities and aggressive interactions. Stoneflies concentrated to 8x natural densities with access to refugia remained longer on snags than a single stonefly without access to refugia. During periods of activity, refugia were defended with larger stoneflies always displacing smaller nymphs. During long periods of inactivity, two and sometimes three nymphs would rest side-by-side sharing the same refuge. Hunger level (starved versus satiated stoneflies), an indirect measure of a predator's response to prey availability, had no significant effect on drift or habitat selection regardless of the presence of refugia. Stonefly predators had a uniform distribution while their prey were clumped. Drift was deliberate and almost always delayed until night, usually at dusk or dawn. An examination of previous research plus the results of this study suggest that non-predatory intra- and interspecific interactions can be an important mechanism causing drift in streams.     

Labiobaetis species of Japan, Taiwan, and Korea, with a new synonym of L. atrebatinus (Eaton 1870) and reerection of the subspecies L. atrebatinus orientalis (Kluge 1983) (Ephemeroptera, Baetidae)

We associated nymphs of Labiobaetis sp. G and Labiobaetis sp. Q from Japan with imagoes reared from nymphs in the field. Labiobaetis sp. G was identified with L. atrebatinus (Eaton 1870) based on characters of the reared male and female imagoes, nymphs, and eggs. We also synonymized a Taiwanese species, L. morus (Chang and Yang 1994), with L. atrebatinus. After further examination of the characters of male imagoes from Japan and Korea and nymphs from Japan and Taiwan, we found them to be correspondent to subspecies L. atrebatinus orientalis (Kluge 1983). Thus, we reerected the subspecific status of L. a. orientalis, although it had been considered not distinguishable from the nominotypical subspecies L. a. atrebatinus. Labiobaetis a. orientalis is distributed in the Russian Far East, Japan, Korea, and Taiwan. We identified Labiobaetis sp. Q with L. tricolor (Tshernova 1928) based on characters of the reared male and female imagoes, nymphs, and eggs. Labiobaetis tricolor was recorded from Japan for the first time.     

Predation and drift of lotic macroinvertebrates during colonization

Summary A field experiment was carried out to determine the effect of an invertebrate predator on the colonization and drift of benthic macroinvertebrates in experimental stream channels. Lotic invertebrates colonized four replicate channels: two controls with no predators, and two channels with low densities (2.8 m^–2) of predatory stonefly nymphs, Doroneuria baumanni (Perlidae). Immigration rates were measured at the inflow of two other channels. Drift rates of invertebrates immigrating to and emigrating from channels were measured daily, and benthic samples were collected every five days. Over a 25-day colonization period, benthic densities of Baetis nymphs and larval Chironomidae were reduced by D. baumanni. Colonization curves were fit with a power function and significantly different colonization rates were indicated for both Baetis and chironomids in predation and control channels. A predator-induced drift response was exhibited by Baetis only and this response was size-dependent. In the presence of D. baumanni, large Baetis drifted more frequently than small nymphs and, correspondingly, small nymphs were more frequent in the benthos. Net predator impacts on invertebrate densities in channel substrates were partitioned into predator-induced drift and prey consumption. These estimates suggest that predator avoidance by Baetis is a prominent mechanism causing density reductions in the presence of predators. Reductions in the density of Chironomidae, however, were attributed to prey consumption only. A rainstorm during the experiment demonstrated that stream flow disruptions can override the influence of predators on benthic invertebrates, at least temporarily, and re-set benthic densities.     

Characterization of benthic microhabitat: An experimental system for aquatic insects

An experimental system used to determine microhabitat current velocity and microhabitat selection by aquatic insects is described. The experimental system includes a microvelocity probe and a hydraulically calibrated artificial substrate. A thermistor velocity probe detects flow velocities to 0.5 m s^–1 near the surface of substrates at locations inhabited by aquatic insects. The artificial substrate was designed to provide two major habitat types, highly turbulent vortex areas and regions with unidirectional, near laminar flow. Substrate calibration and microhabitat characteristics of the substrates are demonstrated. Experimental studies of Simulium sp. location on substrates indicated that while simuliid larvae are characteristic of lotic, erosional habitats, actual microhabitats selected are governed by substantially lower flow velocity.     

Molecular phylogeny evidence of altitudinal distribution and habitat adaptation in Korean Ephemera species (Ephemeroptera: Ephemeridae)

Molecular phylogeny of the four Korean Ephemera species, Ephemera orientalis, E. sachalinensis, E. strigata, and E. separigata, was inferred from 630 bp sequences of the partial mitochondrial cytochrome c oxidase I (COI) gene. Results indicated that mean intraspecific sequence divergences were 0.70%, whereas mean interspecific divergences were 15.75%, and 17 samples were distinguished to four species correctly by COI sequences. The results also demonstrated that four species of Korean Ephemera assembled a monophyletic group with high support in maximum parsimony and maximum likelihood analyses. This Ephemera group was divided into two major clades of E. orientalis–E. sachalinensis and E. strigata–E. separigata. Furthermore, we demonstrated that this phylogeny explained altitudinal and habitat adaptations of Korean Ephemera species. The E. orientalis–E. sachalinensis clade, a widespread and lowland‐adapted mayfly group, retained plesiomorphic traits such as paired stripes on abdominal segments and was regarded as plesiotypic in terms of habitat adaptation, compared to the geographically more limited and upland‐adapted E. strigata–E. separigata clade.     

Small-scale movements of lotic macroinvertebrates with variations in flow

1. The small-scale movements and distribution patterns of invertebrates were observed in an attempt to identify the various mechanisms by which organisms may use flow refugia during flow disturbances. The microdistribution of lotic macroinvertebrates was examined in two replicate, non-circulating laboratory flumes with variations in flow among microhabitat patches (≈ 0.015–0.035 m²). The discharge in one experimental flume was manipulated to mimic spates and alter near-bed flow patterns; the other flume acted as a control. After an initial settling period, the position and behaviour of animals within the flumes was recorded before, during and after a simulated spate. Three species with contrasting flow microhabitat preferences and movement behaviour were examined. 2. At low discharge, the microdistribution of all three study species in flumes was broadly consistent with field observations. In the field, the optimum current speed was lowest for adults of the dytiscid beetle, Oreodytes sanmarkii, and highest for mayfly nymphs, Ephemerella ignita, with nymphs of the stonefly, Leuctra inermis, most abundant at intermediate velocities. In the flumes, O. sanmarkii occurred only in very low velocity areas, L. inermis occurred widely throughout the flumes with highest density in low velocity areas and E. ignita also occurred throughout the flumes, but maximum density was in moderately high velocity areas. 3. Increased discharge did not reduce the total number of individuals in experimental versus control flumes for any of the three species studied, although total numbers did decrease over the observation period in both treatments. Simulated spates resulted in a change in the microdistribution of O. sanmarkii and E. ignita, but not L. inermis, such that numbers were reduced in very high velocity microhabitats and animals accumulated in lower flow areas, analogous to flow refugia. These distributional shifts were attributed to movements of individuals among microhabitats. 4. Both active and passive modes of movement contributed to the accumulation of E. ignita and O. sanmarkii in low flow microhabitats (i.e. flow refugia). Some nymphs of E. ignita actively crawled from high to low flow microhabitats. Both species drifted between microhabitats. Drift entry could be active or passive, whereas regaining the substratum was active: O. sanmarkii swam down and E. ignita altered its body posture to promote sinking.     

How Currents on Different Sides of Substrates in Streams Affect Mechanisms of Benthic Algal Accumulation

Benthic diatom accumulation rates and diversity were monitored for 32 days in five habitats where current conditions were different. After 32 days, diatom abundance in a sheltered habitat where eddies occurred over substrates was over eight times the abundance in a habitat that was exposed directly to the main flow of the stream (about 30 cm/sec). Diatom immigration rates and species richness of diatom taxocenes were negatively related to exposure; whereas, diatom reproduction rates, death rates and relative abundances of the eight dominant diatom species were not related to the small differences in current velocity that occurred. During the 32-day colonization period diatom immigration rates increased, species richness of diatom taxocenes increased and evenness of species abundances decreased. Species composition of diatom taxocenes shifted from a numerical dominance by fast immigrators to dominance by fast reproducers during the 32-day colonization.     

The influence of spatial heterogeneity on the behavior and growth of two herbivorous stream insects

Environmental heterogeneity can affect the behavior of organisms, but the consequences of patchiness for organismal energetics (e.g., growth, fitness) are not well understood. This study demonstrates that spatial heterogeneity can affect the growth of aquatic stream insects in laboratory streams, and reveals the behavioral mechanisms for these effects. In a 2×2 factorial design, I experimentally manipulated resource distribution (homogeneous vs. patchy, with the same overall resource levels) and current velocity (fast vs. slow) to investigate the direct and interactive effects of these factors on the drift behavior and growth of two mobile stream grazers, the mayflies Baetis bicaudatis and Epeorus deceptivus. B. bicaudatis nymphs grew larger in environments with homogeneously distributed resources than in patchy environments, and both species grew larger in fast than slow current environments. Patterns of drift behavior over the course of the study corresponded to observed differences in growth. Both species grew to larger body size in treatments where they drifted more successfully among substrates (fast-current treatments) and where they entered the drift less frequently (fast current for both species, and homogeneous treatments for B. bicaudatis). Overall, these results demonstrate that patchiness can significantly influence both the behavior of aquatic insects and the size to which these insects grow. In the light of previously published relationships between nymphal mayfly body mass and fecundity, these results suggest that patchiness in streams may have important consequences for mayfly populations.     

Use of laboratory studies to develop a dispersal model for Missouri River pallid sturgeon early life intervals

Understanding the drift dynamics of pallid sturgeon (Scaphirhynchus albus) early life intervals is critical to evaluating damming effects on sturgeons. However, studying dispersal behavior is difficult in rivers. In stream tanks, we studied the effect of velocity on dispersal and holding ability, estimated swimming height, and used the data to estimate drift distance of pallid sturgeon. Dispersal was by days 0–10 embryos until fish developed into larvae on day 11 after 200 CTU (daily cumulative temperature units). Embryos in tanks with a mean channel velocity of 30.1 cm s⁻¹ and a side eddy could not hold position in the eddy, so current controlled dispersal. Late embryos (days 6–10 fish) dispersed more passes per hour than early embryos (days 0–5 fish) and held position in side eddies when channel velocities were 17.3 cm s⁻¹ or 21.1 cm s⁻¹. Day and night swim‐up and drift by embryos is an effective adaptation to disperse fish in channel flow and return fish from side eddies to the channel. Early embryos swam <0.50 cm above the bottom and late embryos swam higher (mean, 90 cm). A passive drift model using a near bottom velocity of 32 cm s⁻¹ predicted that embryos dispersing for 11 days in channel flow would travel 304 km. Embryos spawned at Fort Peck Dam, Missouri River, must stop dispersal in <330 km or enter Lake Sakakawea, where survival is likely poor. The model suggests there may be a mismatch between embryo dispersal distance and location of suitable rearing habitat. This situation may be common for pallid sturgeon in dammed rivers.     

Habitat persistence for sedentary organisms in managed rivers: the case for the federally endangered dwarf wedgemussel (Alasmidonta heterodon) in the Delaware River

1. To manage the environmental flow requirements of sedentary taxa, such as mussels and aquatic insects with fixed retreats, we need a measure of habitat availability over a variety of flows (i.e. a measure of persistent habitat). Habitat suitability measures in current environmental flow assessments are measured on a ‘flow by flow’ basis and thus are not appropriate for these taxa. Here, we present a novel measure of persistent habitat suitability for the dwarf wedgemussel (Alasmidonta heterodon), listed as federally endangered in the U.S.A., in three reaches of the Delaware River. 2. We used a two‐dimensional hydrodynamic model to quantify suitable habitat over a range of flows based on modelled depth, velocity, Froude number, shear velocity and shear stress at three scales (individual mussel, mussel bed and reach). Baseline potentially persistent habitat was quantified as the sum of pixels that met all thresholds identified for these variables for flows ≥40 m³ s^?1, and we calculated the loss of persistently suitable habitat by sequentially summing suitable habitat estimates at lower flows. We estimated the proportion of mussel beds exposed at each flow and the amount of change in the size of the mussel bed for one reach. 3. For two reaches, mussel beds occupied areas with lower velocity, shear velocity, shear stress and Froude number than the reach average at all flows. In the third reach, this was true only at higher flows. Together, these results indicate that beds were possible refuge areas from the effects of these hydrological parameters. Two reaches showed an increase in the amount of exposed mussel beds with decreasing flow. 4. Baseline potentially persistent habitat was less than half the areal extent of potentially suitable habitat, and it decreased with decreasing flow. Actually identified beds and modelled persistent habitat showed good spatial overlap, but identified beds occupied only a portion of the total modelled persistent habitat, indicating either that additional suitable habitat is available or the need to improve habitat criteria. At one site, persistent beds (beds where mussels were routinely collected) were located at sites with stable substratum, whereas marginal beds (beds where mussels were infrequently collected or that were lost following a large flood event) were located in scoured areas. 5. Taken together, these model results support a multifaceted approach, which incorporates the effects of low and high flow stressors, to quantify habitat suitability for mussels and other sedentary taxa. Models of persistent habitat can provide a more holistic environmental flow assessment of rivers.     

The effect of current velocity and sediment on the drift of the mayfly Ephemerella subvaria Mcdunnough

SUMMARY. Experiments conducted in an artificial stream showed that significantly more nymphs drifted from an inorganic substrate at a mean current velocity of 28.5 cm s⁻¹ than at 18.5 cm s⁻¹. Drift density, however, was not affected. Disproportionately large numbers of nymphs drifted while current velocities were being increased from 18.5 to 28.5 cm s⁻¹.
Both drift numbers and drift density were greater in turbid water, after the addition of large amounts of inorganic sediment, than under clear-flowing conditions during dark periods but not in the light. The interaction of increasing current velocity and sediment levels resulted in a significantly greater number of drifting nymphs under lighted conditions.
Minor spates which do not seriously disturb the stream bed may initiate significant increases in macroinvertebrate drift.     

Hot rocks or no hot rocks: overnight retreat availability and selection by a diurnal lizard

I used radio telemetry to determine the effects of substrate size and composition on overnight retreat site selection by western fence lizards ( Sceloporus occidentalis). In watersheds of northern California (USA), these lizards occupy two habitat types differing in substrate characteristics: rocky cobble bars found in the dry, active channels of rivers and grassy upland meadows. Rocky substrates, found almost exclusively on cobble bars, provided warmer potential retreat sites than all available retreat sites on meadows during the first 5 h of inactivity. Only cobble and sand substrates provided retreats with temperatures within the preferred daily active range (32–36°C) during the inactive period for these lizards (1900–0900 hours). Females on a cobble bar used rocks as retreats on >90% of nights during the breeding season whereas females on a meadow used wood (>70% of nights) and burrows (>25% of nights). In contrast to females, cobble bar males used rocks significantly less frequently (<70%) and slept in the open air significantly more frequently (25% vs. <1%). Cobble bar females further, showed a significant preference for cobbles 15 cm thick, whereas the rocks used by males did not differ significantly in thickness from those measured in randomly placed transects. Rocks 15 cm thick were the warmest retreats commonly available on this habitat type. Thus, thermal microenvironments available to and chosen by gravid female lizards differ considerably between river and non-river habitats.     

Influences of aquatic macrophytes on invertebrate community structure,guild structure,and microdistribution in streams

The aquatic macrophytes Ranunculus aquatilis and Rorippa nasturtium-aquaticum were transplanted into substrate trays and placed in a stream alongside unvegetated substrate. Macrophytes were observed to have significant effects on 1) invertebrate community structure, 2) guild structure, and 3) microdistribution. 1) Significantly higher taxa richness and community abundances were associated with macrophytes. 2) Significantly higher abundances of shredder, scraper, and predator guilds were associated with macrophytes in fall, and all guilds had higher abundances in macrophytes in spring. However, guild frequency distributions did not differ among habitats except in spring. 3) Enallagma, Gammarus, Gyraulus, Physa, and Pisidium exhibited a strong association with macrophytes, while Hydropsyche, Simulium, Baetis tricaudatus, Glossosoma velona, and Helicopsyche borealis appeared to avoid them. A strong correlation appeared to exist between current velocity preferences of these taxa and their selection or avoidance of vegetated habitat. Thus, the effect of macrophytes in reducing current velocities appeared to be the most important influence on invertebrate microdistribution. However, macrophytes also increase physical heterogeneity and their large surface areas benefit invertebrate community abundances by creating additional living spaces in the water column where none exist above unvegetated substrate.     

Small fish,large river: Surprisingly minimal genetic structure in a dispersal‐limited,habitat specialist fish

Genetic connectivity is expected to be lower in species with limited dispersal ability and a high degree of habitat specialization (intrinsic factors). Also, gene flow is predicted to be limited by habitat conditions such as physical barriers and geographic distance (extrinsic factors). We investigated the effects of distance, intervening pools, and rapids on gene flow in a species, the Tuxedo Darter (Etheostoma lemniscatum), a habitat specialist that is presumed to be dispersal‐limited. We predicted that the interplay between these intrinsic and extrinsic factors would limit dispersal and lead to genetic structure even at the small spatial scale of the species range (a 38.6 km river reach). The simple linear distribution of E. lemniscatum allowed for an ideal test of how these factors acted on gene flow and allowed us to test expectations (e.g., isolation‐by‐distance) of linearly distributed species. Using 20 microsatellites from 163 individuals collected from 18 habitat patches, we observed low levels of genetic structure that were related to geographic distance and rapids, though these factors were not barriers to gene flow. Pools separating habitat patches did not contribute to any observed genetic structure. Overall, E. lemniscatum maintains gene flow across its range and is comprised of a single population. Due to the linear distribution of the species, a stepping‐stone model of dispersal best explains the maintenance of gene flow across its small range. In general, our observation of higher‐than‐expected connectivity likely stems from an adaptation to disperse due to temporally unstable and patchy habitat.     

Predator detection and avoidance by lotic mayfly nymphs of different size

We studied antipredatory responses of lotic mayfly (Baetis) nymphs in a factorial experiment with four levels of fish presence: (1) a freely foraging fish (the European minnow,Phoxinus phoxinus), (2) a constrained fish, (3) water from a fish stream, (4) water from a fishless stream. LargeBaetis nymphs drifted mainly during night-time in treatments involving either the chemical or actual presence of fish, whereas no diel periodicity was observed when the water was not conditioned with fish odour. The response was strongest when the fish was uncaged, which suggests that visual or hydrodynamic cues are needed in addition to chemical ones for an accurate assessment of predation risk. Fish presence had no effect on the drift rates of small nymphs. Instead, they increased their refuge use in the presence of a live fish. Chemical cues alone did not have any effect on the refuge use of any of theBaetis size classes. Our results indicate active drift entry by mayfly nymphs. Because predation pressure is spatially and temporally variable, nymphs must sample the environment in order to locate predator-free areas or areas with low predation risk. Drifting should be the most energy-saving way to do this. To avoid the risk from visually feeding fish, large individuals can sample safely (i.e. enter drift) only at night-time, while the small ones can also do this safely during the day. We suggest that, contrary to some earlier assumptions, mayfly drift is not a fixed prey response. Instead,Baetis nymphs are able to assess the prevailing predation pressure, and they adjust their foraging behaviour accordingly.     

Do pools impede drift dispersal by stream insects?

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