Spatial and temporal variability of macroinvertebrates in spawning and non-spawning habitats during a salmon run in Southeast Alaska

Spawning salmon create patches of disturbance through redd digging which can reduce macroinvertebrate abundance and biomass in spawning habitat. We asked whether displaced invertebrates use non-spawning habitats as refugia in streams. Our study explored how the spatial and temporal distribution of macroinvertebrates changed during a pink salmon (Oncorhynchus gorbuscha) spawning run and compared macroinvertebrates in spawning (riffle) and non-spawning (refugia) habitats in an Alaskan stream. Potential refugia included: pools, stream margins and the hyporheic zone, and we also sampled invertebrate drift. We predicted that macroinvertebrates would decline in riffles and increase in drift and refugia habitats during salmon spawning. We observed a reduction in the density, biomass and taxonomic richness of macroinvertebrates in riffles during spawning. There was no change in pool and margin invertebrate communities, except insect biomass declined in pools during the spawning period. Macroinvertebrate density was greater in the hyporheic zone and macroinvertebrate density and richness increased in the drift during spawning. We observed significant invertebrate declines within spawning habitat; however in non-spawning habitat, there were less pronounced changes in invertebrate density and richness. The results observed may be due to spawning-related disturbances, insect phenology, or other variables. We propose that certain in-stream habitats could be important for the persistence of macroinvertebrates during salmon spawning in a Southeast Alaskan stream.     

Benthic community metabolism in three habitats in an Ozark stream

Benthic community metabolism was measured in three habitats (riffles, runs and pools) during spring (May), summer (July) and fall (October) in the Jacks Fork River, Missouri, using an in situ chamber technique. Net community productivity (NCP) and gross community productivity (GCP) were highest in riffles, lowest in pools and intermediate in runs. Rates of NCP and GCP during spring and fall were similar for both riffles and runs, but NCP and GCP increased significantly during summer in both habitats. Pool substrates were always heterotrophic and exhibited no significant seasonal changes in NCP or GCP. Community respiration (CR) was highest in riffles, intermediate in runs and lowest in pools, but interhabitat differences in CR were generally smaller than for NCP. Rates of CR during spring and fall were similar, but CR increased significantly during summer. Results indicate that the physical conditions associated with each habitat strongly affect benthic community metabolism in this stream and that the relative proportions of these habitats will influence the ratio of living algal:detrital organic matter potentially available for consumers.     

Traits of brown trout prey in relation to habitat characteristics and benthic invertebrate communities

A trait database based on habitat, behavioural and morphological characteristics of invertebrates enabled invertebrates known to have differential vulnerability to salmonid predation to be discriminated. Among all of these traits the most important were associated with invertebrate spatial proximity to salmonids (macrohabitat preference), potential conspicuousness and accessibility (e.g. flow exposure, mobility, tendency to drift, trajectory and agility) and handling efficiency (body flexibility and size). Data on benthos assemblages and brown trout salmo trutta diets collected over two consecutive summers in different macrohabitat types (e.g. pools, runs and riffles) of two Pyrenean streams demonstrated that: (i) decrease in torrential flow characteristics of habitats was associated with changes in invertebrate body shape and movement, and a decrease in body flexibility and tendency to drift; (ii) brown trout tended to feed on the most vulnerable invertebrates (e.g. those living in exposed microhabitats and with a high tendency to drift, agility and aggregation), except in faster flowing habitats. The results suggest that the potential vulnerability of benthic invertebrates increased with increases in flow (torrential) characteristics. Also, the database compiled on traits explained differences among brown trout diets, especially in slow flowing areas.     

Do pools impede drift dispersal by stream insects?

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Habitat overlap between predatory benthic fish and their invertebrate prey in streams: the relative influence of spatial and temporal factors on predation risk

1. The spatial heterogeneity of ecosystems as well as temporal activity patterns of organisms can have far‐reaching effects on predator–prey relationships. We hypothesised that spatiotemporal constraints in mesohabitat use by benthic fish predators would reduce habitat overlap with benthic invertebrates and lead to mesohabitat‐specific predation risks. 2. We analysed the spatiotemporal activity patterns of two small‐bodied benthivorous fishes, gudgeon (Gobio gobio) and stone loach (Barbatula barbatula), and of benthic invertebrates in a small temperate stream during three 24‐h field experiments. By applying a novel method of field video observation, we monitored the spatiotemporal foraging behaviour of the fish in their natural environment. A parallel analysis of invertebrate mesohabitat use by means of small area Hess sampling allowed a direct estimation of habitat overlap at a pool–riffle scale. 3. Gudgeon showed a dominant spatial activity pattern preferring pools at all times of day, whereas stone loach used both mesohabitats but with a distinct temporal (nocturnal) activity pattern. The patterns of residence were not identical with those of active foraging. Invertebrate community composition differed significantly between mesohabitats but not between times of day. More than half of the total dissimilarity between pools and riffles was accounted for by six invertebrate taxa. Five of these were subject to higher fish predation in pools than in riffles. The total prey consumption of the two fish species together in pools was about three times as high as in riffles. Trophic niche breadth of stone loach and thus its predation range was broader than that of gudgeon. 4. These results indicate that the potential predation risk for stream invertebrates depends on the combination of spatial and temporal patterns of both predator and prey. Given the distinct differences in predation risk found between pools and riffles, we conclude that spatial heterogeneity at the mesohabitat scale can influence mechanisms and consequences of selective predation. We also suggest that the analysis of spatiotemporal predator–prey relationships should not be based on the premise that the main residence habitat and active foraging habitat of a predator are identical.     

Comparisons of benthic invertebrates between riffles and pools

Benthic species assemblages in upstream and downstream ends of riffles and in pools were investigated seasonally in the first five orders of an alluvial gravel stream with distinct pool and riffle channel form. Riffles comprised < 10% of stream area and were separated by pools with extensive bedrock substrate (ca. 15–85% of total surface area) which was scoured during periodic high flow. Virtually all taxa were more abundant in riffles than in pools, except chironomids which were more equally distributed. Inconsistent results were obtained for upstream-downstream comparisons within riffles. Intermittent headwater reaches (orders 1 & 2) which supported half as many taxa retained this pattern during periods of flow, although riffles at these sites were dry from mid-June to mid-November. Pools which contained more gravel, indicating less disturbance during high flow, had a richer assemblage of benthic species than other pools. Many invertebrates in pools may have been there as a result of drift from their preferred riffle habitats, but the presence of gravel in the pools indicates less intense flow disturbance during floods, provides protection from the mild scouring that does occur during floods, and provides refugia from predators.     

Local Movement as a Measure of Habitat Quality in Stream Salmonids

Habitat assessments are often based on the premise that spatial variation in population density arises from, and accurately reflects, underlying differences in quality among habitats. Nonetheless, this premise has been criticized on both theoretical and empirical grounds. Habitat quality perhaps is best evaluated by examining behavioural processes which directly influence habitat use. We present an approach based on the assumption that measures of local movement, such as habitat-specific immigration and loss rates, provide useful indicators of habitat quality. A dynamic turnover model was used in conjunction with mark-recapture techniques to estimate movement parameters for brook charr, Salvelinus fontinalis, and Atlantic salmon, Salmo salar, in different stream habitats during the summer. Immigration and loss rates were derived from mark-recapture experiments covering short periods of time (6 days). Movement-based rankings of habitat quality for both charr (pools glides > riffles) and salmon (riffles > glides > pools) were in agreement with results from earlier studies. Over evaluation periods of up to 65 days, observed abundances were highly variable in time and fluctuated about the equilibrium abundances calculated from movement parameters in the short-term experiments, suggesting that movement-based parameters may be more stable than measures of abundance for evaluating salmonid habitat. Because the approach based on movement behaviour focuses on immigration and loss, two processes that directly generate density differentials between habitats, it provides a more reliable mechanistic basis for understanding habitat selection than do traditional approaches based on density–quality relationships.     

Identifying the scales of variability in stream macroinvertebrate abundance, functional composition and assemblage structure

1. Many natural ecosystems are heterogeneous at scales ranging from microhabitats to landscapes. Running waters are no exception in this regard, and their environmental heterogeneity is reflected in the distribution and abundance of stream organisms across multiple spatial scales. 2. We studied patchiness in benthic macroinvertebrate abundance and functional feeding group (FFG) composition at three spatial scales in a boreal river system. Our sampling design incorporated a set of fully nested scales, with three tributaries, two stream sections (orders) within each tributary, three riffles within each section and ten benthic samples in each riffle. 3. According to nested anova s, most of the variation in total macroinvertebrate abundance, abundances of FFGs, and number of taxa was accounted for by the among‐riffle and among‐sample scales. Such small‐scale variability reflected similar patterns of variation in in‐stream variables (moss cover, particle size, current velocity and depth). Scraper abundance, however, varied most at the scale of stream sections, probably mirroring variation in canopy cover. 4. Tributaries and stream sections within tributaries differed significantly in the structure and FFG composition of the macroinvertebrate assemblages. Furthermore, riffles in headwater (second order) sections were more variable than those in higher order (third order) sections. 5. Stream biomonitoring programs should consider this kind of scale‐dependent variability in assemblage characteristics because: (i) small‐scale variability in abundance suggests that a few replicate samples are not enough to capture macroinvertebrate assemblage variability present at a site, and (ii) riffles from the same stream may support widely differing benthic assemblages.     

Seasonal and diel periodicity in the drift of aquatic insects in a subtropical Florida stream

A study of insect drift was conducted in a small, subtropical Florida stream from December 1971 to December 1972 to describe the seasonal and diel periodicity and to determine factors influencing behavioural drift. Paired samples of 2 h duration beginning 15 min after sunset were taken biweekly, and hourly collections over a 24-h period were made quarterly. Benthic invertebrates were collected on each date from three habitats (riffle, pool and aquatic vegetation) and temperature, dissolved oxygen and current velocity were measured. Drift rates ranged from 100 to 2125 organisms/m². h (0·03 to 0·49 organisms/m³) and were greatest in winter and early spring; minimal rates occurred in the summer months. The following six taxa, in order of relative abundance, comprised 87% of the drift: Baetis intercalaris, Cheumatopsyche sp., Stenonema exiguum, Chironomidae, Stenelmis fuscata and Simulium sp. Total drift showed no significant correlation with temperature, dissolved oxygen or mean benthic abundance and only slight correlation with current velocity (r=0·34). Stepwise, multiple regression analyses indicated that riffle density and mean size of drifting organisms were important factors influencing the drift rates of B. intercalaris (R=0·67) and S. exiguum (R=0·82); mean size, riffle density and water temperature influenced the drift of Cheumatopsyche sp. (R=0·78). The other taxa of drifting insects showed no significant correlation with the variables measured. Diel (24 h) studies of the major taxa showed marked differences in the periodicity, both within and between taxa, indicating the need for long-term studies with frequent sample intervals in subtropical habitats. A new drift pattern for the family Chironomidae, alternans type, was observed for late instars of Polypedilum halterale.     

Mesohabitat use by bullhead (Cottus gobio)

Habitat composition and connectivity within a stream vary with changing flows but the influence of changing flow on habitat use by fish is not well understood. Meso- and microhabitat surveys were used to investigate habitat use by bullhead (Cottus gobio Linnaeus) in response to discharge variation in a small tributary of the Upper Severn, England. Mesohabitat mapping surveys were carried out over a range of summer flows (0.016–0.216 m³ s⁻¹) and were coupled with direct underwater observations (snorkelling) of fish location. Five mesohabitat types—glides, runs, riffles, chutes and pools—were present in the reach at all flows surveyed and ‘backwaters’ were found at three flows. The macro-morphology of the reach comprised six riffle–pool sequences divided into 27 mesohabitats with the maximum diversity (23 mesohabitats) at intermediate flows (Q ₄₃) and only 15 mesohabitats at Q ₉₅. Despite low numbers of fish (N = 78), bullhead displayed a strong association (51% of the fish) with glides—relatively deep habitats having high rates of velocity increase with flow. However, 54% of the fish were observed in two large, persistent mesohabitats, a glide (34%) and a pool (20%), both located below a faster flowing mesohabitat. Habitat use curves based upon micro-habitat data showed bullhead favoured low velocities (<0.30 m s⁻¹), depths less than 0.30 m and a cobble substratum. This study illustrates the value of cross-scale investigations in linking fish ecology, flow and physical habitat variability and suggests mesohabitat size, persistence and arrangement may influence fish distribution.     

Linking stream habitats and spider distribution: spatial variations in trophic transfer across a forest–stream boundary

In headwater streams, many aquatic insects rely on terrestrial detritus, while their emergence from streams often subsidizes riparian generalist predators. However, spatial variations in such reciprocal trophic linkages remain poorly understood. The present study, conducted in a northern Japanese stream and the surrounding forest, showed that pool–riffle structure brought about heterogeneous distributions of detritus deposits and benthic aquatic insects. The resulting variations in aquatic insect emergence influenced the distributions of riparian web-building spiders. Pools with slow current stored greater amounts of detritus than riffles, allowing more benthic aquatic insects to develop in pools. The greater larval biomass in pools and greater tendency for riffle insects to drift into pools at metamorphosis resulted in an emergence rate of aquatic insects from pools that was some four to five times greater than from riffles. In the riparian forest, web-building spiders (Tetragnathidae and Linyphiidae) were distributed in accordance with the emergence rates of aquatic insects, upon which both spider groups heavily depended. Consequently, the riparian strips bordering pools had a density of tetragnathid spiders that was twice as high as that of the riparian strips adjacent to riffles. Moreover, although limitations of vegetation structure prevented the aggregation of linyphiid spiders around pools, linyphiid density normalized by shrub density was higher in habitats adjacent to pools than those adjacent to riffles. The results indicated that stream geomorphology, which affects the storage of terrestrial organic material and the export of such material to riparian forests via aquatic insect emergence, plays a role in determining the strength of terrestrial–aquatic linkages in headwater ecosystems.     

Influence of morphohydraulic habitat structure on invertebrate communities (Ephemeroptera,Plecoptera and Trichoptera)

Fluvial geomorphology proposes the methodology of cognition and assessment of the riverine landscape and points to the possibilities of exploitation of its results in hydrobiological research. Habitat structure of two reaches of the Drietomica brook (Biele Karpaty Mts, Slovakia) was assesed at level of morphological and morphohydraulic units in the sense of the River Morphology Hierarchical Classification (RMHC)). Physical habitats were described by flow hydraulics and substrate properties as directly measured variables (current velocity, depth, substrate size) and related variables (flow type, Froude and Reynolds numbers). According to the shear stress (expressed by Fr and Re), the morphological units were divided into two main groups — with low shear stress — pools, glides, edgewaters, bar nooks and bars; with high shear stress — riffles, runs, rapids and scours; characterized also by different Ephemeroptera, Plecoptera and Trichoptera (EPT) communities. The EPT communities were analyzed in relation to the morphological, hydraulic and substrate characteristics of the stream channel. The main environmental gradient responsible for the variation in EPT fauna was found using Principal Component Analysis and was related to gradient of flow in term of current velocity and other hydraulic attributes covered by Fr and Re numbers. The EPT communities (by means of abundance, feeding types, current, microhabitat and zonation preferences) showed preferences for different morphological units, flow type and current velocity. Depth and substrate grain size showed only weak relation to EPT communities.     

Hydraulic microhabitats and the distribution of macroinvertebrate assemblages in riffles

1. The influence of hydraulic conditions on the spatial distribution of macroinvertebrate assemblages was investigated in three riffles in a perennial Australian river. 2. Velocity, depth and variability of substrate roughness were measured at each of 56 macroinvertebrate sampling locations. Complex hydraulic variables (roughness Reynolds number, shear velocity, Froude number) were calculated from combinations of two or three of the directly measured variables. The biological significance of directly measured and complex hydraulic variables was determined by a combination of univariate and multivariate statistical procedures. 3. Macroinvertebrate abundance, number of taxa and community composition were significantly different between the identified roughness Reynolds number, Froude number, velocity and shear velocity microhabitats throughout the studied riffles. 4. Regression analysis showed macroinvertebrate abundance and number of taxa were negatively related to roughness Reynolds number, shear velocity, velocity and Froude number. Depth was negatively related to abundance. In general, the majority of the macroinvertebrate community preferred the areas of riffles with the lowest near‐bed turbulence. 5. Roughness Reynolds number explained more of the spatial variation in invertebrate abundance, number of taxa and community composition than the other hydraulic variables, either directly measured or calculated. Of the directly measured variables, velocity had the greatest explanatory power, which was marginally less than roughness Reynolds number and shear velocity. 6. This study demonstrated that small‐scale differences in hydraulic conditions created by combinations of velocity, depth and substrate roughness have an important role in the spatial distribution of macroinvertebrate assemblages in riffle habitats.     

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     

Experimentally induced foraging mode shift by sympatric charrs in a Japanese mountain stream

Foraging mode shifts may allow animals flexibility to adaptto a variety of ecological conditions. Theory holds that ectothermssuch as fishes should shift from ambush to active search modeswhen prey density declines, to maintain a minimum encounterrate. Salvelinus malma (Dolly Varden) sympatric withS. leucomaenis(white-spottedchirr) in a northern Japan mountain stream were observed toshift from ambushing drifting invertebrates at fixed focal pointsto picking benthic invertebrates from the substrate along circuitousroutes throughout pools, as drift declined throughout summer.Experimentally reducing drift in two pools during early summerwhen all charr were drift foragers caused subordinate malmato make marked shifts from drift to benthic foraging within1 h, whereas nearly all leucomaenis remained drift foragers.Some charr of both species responded to reduced drift by emigratingfrom pools. Combined results of three replicate experimentsrevealed that malma shifted to benthic foraging when their rateof drift forays was reduced below a specific threshold, therebyupholding predictions of the theory. Adaptive foraging modeshirts may promote coexistence of these congeneric charrs inreaches of Japanese mountain streams     

Relationships between mesohabitats, ichthyological communities and IBI metrics adapted to a European river basin (The Meuse, Belgium)

We investigated the influence of mesohabitats on fish communities and on attributes of a multimetric index of fish integrity in the River Meuse basin. Three consecutive 150 m sectors in a Meuse tributary (Ourthe, Belgium), each divided in two or three sub-sections presenting various percentages of mesohabitats (riffles, runs and pools), were sampled by electrofishing. In each sub-section, relative and absolute biomasses of each fish species were estimated. The presence of limnophilic cyprinids was inversely correlated (r ² = 0.70 and 0.56 for absolute and relative biomass, respectively) with the percentage of riffles. Salmonids preferred runs and their absolute biomass was highly dependent (r ² = 0.71) on the proportion of this mesohabitat, whereas biomass of limnophilic cyprinids was highly correlated (r ² = 0.75 and r ² = 0.82 for absolute and relative biomass, respectively) with pools. A positive correlation (r ² = 0.58) was established between relative biomass of predators and the percentage of this mesohabitat. An IBI was calculated for the three sectors on the basis of results from the entire Meuse catchment. Scores of most metrics showed low variation among sectors but values of two metrics (% of individuals as tolerant, % of individuals as ubiquitous spawners) were greatest in sector 2, where pools predominated. Further, the lowest IBI score (51/65, integrity class: fair to good) was recorded in sector 2 where pools dominated, while sector 1 (where runs dominated) obtained the highest score (63/65, integrity class: excellent). Sector 3 which has a balanced proportion of riffles and runs obtained an intermediate score (57/65). Considering the response of IBI to the natural variation of mesohabitat proportions, it appears that an accurate sampling requires the prospection of a variety of mesohabitats (with a majority of runs) for the evaluation of river quality.     

Effect of riparian land use on contributions of terrestrial invertebrates to streams

Since terrestrial invertebrates are often consumed by stream fishes, land-use practices that influence the input of terrestrial invertebrates to streams are predicted to have consequences for fish production. We studied the effect of riparian land-use regime on terrestrial invertebrate inputs by estimating the biomass, abundance and taxonomic richness of terrestrial invertebrate drift from 15 streams draining catchments with three different riparian land-use regimes and vegetation types: intensive grazing — exotic pasture grasses (4 streams), extensive grazing — native tussock grasses (6 streams), reserve — native forest (5 streams). Terrestrial invertebrate drift was sampled from replicated stream reaches enclosed by two 1 mm mesh drift nets that spanned the entire channel. The mean biomass of terrestrial invertebrates that entered tussock grassland (12 mg ash-free dry mass m^–2 d^–1) and forest streams (6 mg AFDM m^–2 d^–1) was not significantly different (p > 0.05). However, biomass estimated for tussock grassland and forest streams was significantly higher than biomass that entered pasture streams (1 mg AFDM m^–2 d^–1). Mean abundance and richness of drifting terrestrial invertebrates was not significantly different among land-use types. Winged insects contributed more biomass than wingless invertebrates to both pasture and tussock grassland streams. Winged and wingless invertebrates contributed equally to biomass entering forest streams. Land use was a useful variable explaining landscape-level patterns of terrestrial invertebrate input for New Zealand streams. Evidence from this study suggests that riparian land-use regime will have important influences on the availability of terrestrial invertebrates to stream fishes.     

Trophic interception: how a boundary-foraging organism influences cross-ecosystem fluxes

The rate at which subsidies move between habitats is a function not only of those factors that facilitate such transfers, but also of factors that limit or restrict the movement of subsidies. The interruption or redirection of subsidies by organisms foraging at the boundary between habitats, or trophic interception, has the potential to substantially restructure the food webs of recipient habitats. In this study we describe how a waterstrider, Aquarius remigis , limits the transfer of a subsidy across the land-water boundary. Prey interception varied with the type of stream habitat; on average, waterstriders in isolated pools intercepted 71.8% of experimental prey inputs of all sizes compared with 21.5% in connected pools and 0.8% in riffles. Across all stream habitat types, waterstriders consumed 43.2% of the experimental inputs of the smallest prey used, the prey size class most similar to natural inputs in our study area. We estimate that foraging waterstriders may intercept 0.3–1.2 kg of terrestrial prey subsidy for every 100 m of stream channel during three summer months, resources that could otherwise support 13–58 young-of-the-year salmonids. In controlled trials, waterstriders significantly altered the amount of terrestrial prey directly consumed by fish, while fish also altered waterstrider interception of those prey. Interestingly, when waterstriders and fish were present together, more terrestrial prey were lost to the streambed than when either or both of these consumers were absent, making this resource available to benthic detritivores, and facilitating the direct incorporation of terrestrial nutrients into stream detrital webs. Overall, we demonstrate that organisms that forage at habitat boundaries can control the quantity and quality of subsidies arriving in recipient habitats, potentially altering food web structures within those habitats.     

山地河流浅滩深潭生境大型底栖动物群落比较研究——以重庆开县东河为例

浅滩和深潭是山地河流中常见的河流生境结构。2011年7月,在重庆开县东河上游双河口-杉木桥河段,选择21个浅滩和深潭,调查大型底栖动物,研究影响不同生境中底栖动物组成、分布和多样性的生态机理。结果表明:调查河段浅滩和深潭中大型底栖动物分别为31种和24种,密度分别为450.62 个/m²和86.24 个/m²,生物量分别为2.88 g/m²和0.55 g/m²。浅滩有指示种11种,即纹石蛾(Hydropsyche sp.)、假蜉(Iron sp.)、假二翅蜉(Pseudocloeon sp.)、舌石蛾(Glossosoma sp.)、高翔蜉(Epeorus sp.1)、背刺蜉(Notacanthurus sp.)、Heterocloeon sp、锯形蜉(Serratella sp.)、朝大蚊(Antocha sp.)、等蜉(Isonychia sp.)、溪颏蜉(Rhithrogena sp.)。深潭指示种仅蜉蝣(Ephemera sp.)和黑大蚊(Hexatoma sp.)两种。刮食者为两类生境的优势功能摄食类群。浅滩中滤食者和刮食者比例显著高于深潭,而收集者和捕食者显著低于深潭。两类生境中大型底栖动物群落结构差异显著。浅滩中大型底栖动物的密度、生物量、丰富度指数、Shannon-Wiener 指数、改进的Shannon-Wiener指数均明显高于深潭。受地貌形态、水力特征和冲淤变化规律影响的生境稳定性和异质性差异是导致大型底栖动物群落差异的主要原因。