Riffle-pool geomorphology disrupts longitudinal patterns of stream benthos

An Ozark Plateau stream was studied to determine the influence of distinct pool and riffle geomorphology on the longitudinal zonation of macroinvertebrate species assemblages and functional group classification. All study sites were dominated by alluvial pool and riffle channel form and the first two orders became intermittent during summer months. Nine benthos samples were collected seasonally from riffles and pools at each of five sites using a vacuum benthos sampler. Diel temperature pulse and coarse particulate organic matter (CPOM) were measured at each site also. Water temperature was most variable in second order, and CPOM on riffles was not more abundant in upstream reaches. Annual average density and biomass of invertebrates were highest in third and fourth orders, respectively. Diversity was depressed in the intermittent headwaters sites. Macroinvertebrate functional groups did not exhibit strong longitudinal trends as predicted by the river continuum model, with species assemblages apparently more strongly affected by the segment-level physical template, although shredders were more abundant in the headwaters during fall and winter. This study indicates that a reach-level perspective based on channel form is a necessary complement to holistic stream ecosystem models, especially in alluvial gravel streams.     

Effects of habitat structure on habitat use by Gammarus pulex in artificial streams

1. The influence of coarse substratum and flow, coarse substratum and food, and predation risk and flow on habitat use by Gammarus pulex was studied in three experiments conducted in artificial stream channels. Each stream channel consisted of a riffle and pool habitat.
2. Location of coarse substrata and food was manipulated by placing cobbles (coarse substratum) and leaf packs (food) in different habitats. Predation risk was varied by running experiments in the presence and absence of sculpins ( Cottus gobio ), and flow was varied by pumping water with one or two pumps.
3. In all experiments Gammarus were most abundant in pools but placement of cobbles in riffles increased use of the latter. An even greater percentage of Gammarus used riffles if leaf packs were also placed there. Decreased discharge and the presence of sculpins ( Cottus gobio ) also caused Gammarus to increase use of riffles. These data indicate that Gammarus is able to evaluate differences in habitat quality and respond accordingly.     

Effects of habitat structure on habitat use by Gammarus pulex in artificial streams

1. The influence of coarse substratum and flow, coarse substratum and food, and predation risk and flow on habitat use by Gammarus pulex was studied in three experiments conducted in artificial stream channels. Each stream channel consisted of a riffle and pool habitat.
2. Location of coarse substrata and food was manipulated by placing cobbles (coarse substratum) and leaf packs (food) in different habitats. Predation risk was varied by running experiments in the presence and absence of sculpins ( Cottus gobio ), and flow was varied by pumping water with one or two pumps.
3. In all experiments Gammarus were most abundant in pools but placement of cobbles in riffles increased use of the latter. An even greater percentage of Gammarus used riffles if leaf packs were also placed there. Decreased discharge and the presence of sculpins ( Cottus gobio ) also caused Gammarus to increase use of riffles. These data indicate that Gammarus is able to evaluate differences in habitat quality and respond accordingly.     

Effects of travertine and flow on leaf retention in Fossil Creek,Arizona

Leaf retention is important in transferring energy from riparian trees to stream food webs. Retention increases with geomorphic complexity such as substrate coarseness, sinuosity, and the presence of debris dams. High discharge can reduce retention, particularly when streams lack physical trapping features. Travertine formations, caused by calcium carbonate deposition, can alter stream morphology. To date, however, we know of no study testing the effect of travertine on leaf retention. This study capitalized on a river restoration project in Fossil Creek, Arizona, where water was returned to the channel after a century of diversion. We examined how the fixed factors Flow (before and after restoration) and Morphology (travertine and riffle-pool sites) affected leaf retention. Leaf retention was higher in sites where travertine forms barriers across the river, relative to sites with riffle-pool morphology. Most leaves retained in travertine reaches were concentrated at the bottom of pools formed between dams. Although flow restoration did not alter retention rates across all sites, it diminished them at travertine sites, indicating an interaction between stream flow and morphology. We conclude that stream complexity and leaf retention are enhanced by travertine deposition but that high discharge can reduce the retentive capacity of in-stream structures. Handling editor: Darren Ryder     

Food,microhabitat, or both? Macroinvertebrate use of leaf accumulations in a montane stream

1. Leaf packs in streams may serve as food and substrate to many macroinvertebrates, but the relative importance of these two functions has not been disentangled. To test the hypothesis that leaf packs are colonized primarily for their food value rather than as microhabitat, the colonization of leaf packs of red alder and of polyester cloth in a natural stream was compared. 2. Species of shredders showed large differences in the colonization of the two types of leaf pack with almost no use of artificial leaf packs. Non-shredders were also more abundant on natural leaf packs, however, they colonized artificial leaf packs six to eighty times proportionally more than shredders. 3. The effect of different leaf types was virtually eliminated for non-shredders when the amount of fine parriculate detritus (food for many non-shredders) trapped in the leaf pack was added to the analysis as a covariate. Therefore, to non-shredders, leaf types differed only in the retention of fine particulate organic matter. 4. Comparisons of the use of both kinds of leaf packs in riffles versus pools revealed that significantly fewer animals colonized pool leaf packs. 5. These results suggest that food value, and not microhabitat, is the primary determinant of leaf pack use for most shredders and non-shredders.     

Flow-substrate interactions create and mediate leaf litter resource patches in streams

1. The roles that streambed geometry, channel morphology, and water velocity play in the retention and subsequent breakdown of leaf litter in small streams were examined by conducting a series of field and laboratory experiments. 2. In the first experiment, conditioned red alder (Alnus rubra Bongard) leaves were released individually in three riffles and three pools in a second‐order stream. The transport distance of each leaf was measured. Several channel and streambed variables were measured at each leaf settlement location and compared with a similar number of measurements taken at regular intervals along streambed transects (‘reference locations’). Channel features (such as water depth) and substrate variables (including stone height, stone height‐to‐width ratio, and relative protrusion) were the most important factors in leaf retention. 3. In the second experiment, the role of settlement location and reach type in determining the rate of leaf litter breakdown was examined by placing individual conditioned red alder leaves in exposed and sheltered locations (on the upper and lower edges of the upstream face of streambed stones, respectively) in riffle and pool habitats. After 10 days, percent mass remaining of each leaf was measured. Generally, leaves broke down faster in pools than in riffles. However, the role of exposure in breakdown rate differed between reach types (exposed pool > sheltered pool > sheltered riffle > exposed riffle). 4. In the third experiment, the importance of substrate geometry on leaf litter retention was examined by individually releasing artificial leaves upstream of a series of substrate models of varying shape. Substrates with high‐angle upstream faces (were vertical or close to vertical), and that had high aspect ratios (were tall relative to their width), retained leaves more effectively. 5. These results show that streambed morphology is an important factor in leaf litter retention and breakdown. Interactions between substrate and flow characteristics lead to the creation of detrital resource patchiness, and may partition leaf litter inputs between riffles and pools in streams at baseflow conditions.     

凉水和帽儿山地区低级溪流生境和水质状况

为研究不同植被景观土地利用下低级溪流的生境状况和水质差异,对凉水国家级自然保护区红松原始林内和帽儿山国家森林公园境内天然次生林植被和农田背景下各3条溪流生境和水质状况进行调查。对溪流生物特性（悬浮藻、附着藻）和理化性质（温度、混浊度、溶解氧（DO）、pH、NH4^＋-N、NO3^-N、PO4^3-P、总氮（TN）和总磷（11P））以及细小颗粒有机物质（FPOM）和粗大有机物质（CPOM））进行测定。研究结果表明,凉水地区原始林溪流的生境状况好于帽儿山地区的次生林,帽儿山地区农田溪流生境最差。原始林溪流具有稳定的溪底生物生活基质,稳水区和急流区均匀分布,稳水区尺度变化大,受淀积物沉降干扰小,河道较弯曲,河岸稳定,河岸植被覆盖度高;次生林溪流以急流区为主,稳定基质相对较差,并受到一定程度的淀积物沉降干扰,河岸尚稳定,有一定程度的人为干扰;农田溪流基质不稳定,受到强烈淀积物沉降影响,渠道化严重,河岸带植被严重破坏。3种景观背景下溪流总磷（11P）、溶解氧（DO）、混浊度、温度、氮磷比值（N／P）（P〈0．05）存在显著差异。原始林溪流NH4^＋、DO、TP、TN、悬浮藻浓度和pH较高、附着藻数量较多,温度较低、FPOM和CPOM的数量较少;次生林溪流的NO3^--N、N／P和TDIN较高。 浊度较低;农田溪流浊度、温度、PO4^3--P较高,DO和pH较低,附着藻数量较少。景观尺度上的土地利用对溪流生境具有深刻的影响,同时决定溪流的水质状况。     

Habitat structure and invertebrate assemblages on stream stones: A multivariate view from the riffle

Abstract Highly structured habitats, those with complex or heterogeneous physical structure, commonly contain more species than simply structured ones. However, tests of this relationship have been hampered by habitat-specific definitions and measurements of habitat structure and, in stream studies, by insufficient information on spatial variation in faunal abundances. We sampled the fauna of 90 stones collected from an order 4 section of the Taggerty and Steavenson Rivers, southeastern Australia. The spatially nested sampling design encompassed three spatial scales (sites within the same stream order, riffles at the same site and groups of stones within the same riffle) but no temporal replication, as we were only able to sample once. Habitat structure of stones was quantified by measuring the amount and type of epilithon on stones, together with stone shapes, sizes, textures and surface complexities. For the latter two attributes, we used a new method, involving image analysis, which can be used to quantify habitat structure in comparable ways in other systems. Species richness (S) was related to some measures of habitat structure, being higher on stones with a rough texture and weakly related to the amount of epilithon present. Total numbers of individuals (N) were also higher on rough stones, and faunal composition (as quantified by hybrid multidimensional scaling) was related to abundance of epilithon. However, flow environments, as quantified by water velocities and depths, were also related to faunal composition. Riffles varied in both faunal densities and near-bed flow environment, but not in any consistent way. Such variability means that individual riffles cannot be reasonably used to ‘represent’ stream sections or orders, an assumption commonly made in stream studies. Neglect of such issues has also resulted in poor conceptual integration between large- and small-scale studies in stream ecology.     

Influence of habitat structure and fish density on Atlantic salmon Salmo salar L. territorial behaviour

A laboratory stream experiment was performed to test the influence of young‐of‐ the‐year Atlantic salmon Salmo Salar density on habitat selection ( i.e . pools and riffles) and the combined effect of habitat and density on territoriality. Pools were preferred at low and medium fish density. At higher density, the saturation of the riffles led to an equal fish repartition in both habitats. Aggressive interactions increased according to fish density, and the level of aggression was significantly higher in the pools than in the riffles.     

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.     

Spatial scale and the diversity of macroinvertebrates in a Neotropical catchment

1. Lotic ecosystems can be studied on several spatial scales, and usually show high heterogeneity at all of them in terms of biological and environmental characteristics. Understanding and predicting the taxonomic composition of biological communities is challenging and compounded by the problem of scale. Additive diversity partitioning is a tool that can show the diversity that occurs at different scales. 2. We evaluated the spatial distribution of benthic macroinvertebrates in a tropical headwater catchment (S.E. Brazil) during the dry season and compared alpha and beta diversities at the scales of stream segments, reaches, riffles and microhabitats (substratum types: gravels, stones and leaf litter). We used family richness as our estimate of diversity. Sampling was hierarchical, and included three stream segments, two stream reaches per segment, three riffles per reach, three microhabitats per riffle and three Surber sample units per microhabitat. 3. Classification analysis of the 53 families found revealed groups formed in terms of stream segment and microhabitat, but not in terms of stream reaches and riffles. Separate partition analyses for each microhabitat showed that litter supported lower alpha diversity (28%) than did stones (36%) or gravel (42%). In all cases, alpha diversity at the microhabitat scale was lower than expected under a null model that assumed no aggregation of the fauna. 4. Beta diversity among patches of the microhabitats in riffles depended on substratum type. It was lower than expected in litter, similar in stone and higher in gravel. Beta diversities among riffles and among reaches were as expected under the null model. On the other hand, beta diversity observed was higher than expected at the scale of stream segments for all microhabitat types. 5. We conclude that efficient diversity inventories should concentrate sampling in different microhabitats and stream sites. In the present study, sampling restricted to stream segments and substratum types (i.e. excluding riffles and stream reaches) would produce around 75% of all observed families using 17% of the sampling effort employed. This finding indicates that intensive sampling (many riffles and reaches) in few stream segments does not result in efficient assessment of diversity in a region.     

Life history studies of anisocentropus kirramus neboiss (trichoptera: Calamoceratidae) in a tropical Australian rainforest stream

Eggs and larvae of Anisocentropus kirramus were collected from leaf packs in riffles and pools in a small upland rainforest stream in tropical Queensland. Adults were collected in floating emergence traps. Egg masses contained 80–100 eggs. None developed in water at 12–15°C but at 22–25°C larvae hatched in 3–10 days. There were five larval instars and complete development appeared to take several months. Instars II‐V were present in all nine months sampled; instar I was present in all but one month. Adults emerged in all ten months sampled, but there was clear seasonality with peak emergence in the summer. The sex ratio of adults was 1:1.     

Eucalypt leaf decomposition in an intermittent stream in south-eastern Australia

Eucalypt leaf packs were placed at two sites in an intermittent stream during summer to examine the hypothesis that terrestrially-exposed leaf litter accumulates a richer microbial flora than submerged leaves — a phenomenon observed in Canadian temporary vernal pools. This did not occur; during the experiment, microbial biomass (as ATP) rose steadily on submerged leaves but remained low on terrestrially-exposed leaves. Densities of most functional feeding groups on the submerged leaves increased with time. Scrapers appeared to be more important than shredders in eucalypt leaf breakdown at both sites.     

Submerged undercut banks as macroinvertebrate habitat in a subalpine meadow stream

We compared macroinvertebrate samples from submerged undercut banks with samples from rifles and pools in a low-gradient-meadow reach of a small stream on the Snowy Range, southeastern Wyoming, from July to September 1985. Macroinvertebrates collected from submerged undercut banks were similar to those collected from riffles and pools, but densities varied among the three habitat types. In July Ephemeroptera, Plecoptera, Trichoptera, Coleoptera, and Diptera, as well as Hydracarina, were all substantially more numerous in samples from undercut banks than from riffles or pools. By contrast in September little difference was observed in the abundance of macroinvertebrates in samples from submerged undercut banks, riffles and pools. While forming only 8.5% of the area of aquatic habitat within the study reach submerged undercut banks were estimated to contain 44% of the aquatic insects in July and 30% in August. Further, submerged undercut banks are macroinvertebrate habitat in this subalpine meadow stream, contributing to overall macroinvertebrate abundance and probably an additional food source for trout.     

Hydrologic versus geomorphic limitation on CPOM storage in stream ecosystems

1. Stream ecosystems are the products of interactions between hydrology, geomorphology and ecology, but examining all three components simultaneously is difficult and rarely attempted. Frequently, either geomorphology or hydrology is treated as invariable or static. 2. To examine the validity of treating either hydrology or geomorphology as static, we studied the individual and combined effects of hydrology and channel geomorphology on coarse particulate organic matter (CPOM) storage. Using data from an experimental leaf release in a hydrologically regulated stream we created a simple numerical model. This allowed us to quantify the relative influence of CPOM trapping and CPOM retention on total long‐term CPOM storage under variable regimes of flood frequency and geomorphic structure. 3. CPOM storage is a function of supply, flood frequency and the type and frequency of in‐stream structures. In‐stream structures perform two distinct functions, trapping and retention, whose relative importance in leaf storage changes with stream hydrology. Trapping is more important for CPOM storage in streams with few floods, while retention is more important in streams with frequent floods. Different structures (e.g. boulders, large wood, small wood) perform these functions at different efficiencies. We found that large wood trapped two to three times more leaves than the bank, but that the bank retained leaves two to three times more efficiently. 4. A modelled channel with five times the amount of large wood as the study channel (a ‘wood restoration’) initially stored 14% more leaves than the modelled ‘natural’ channel. After six floods, however, the modelled wood restoration channel stored 50% less CPOM than the natural channel as the large wood had high trapping but poor retention. The modelled natural channel contained structures that could both trap and retain. Thus, as different structures performed different functions, the structural complexity buffered the stream allochthonous energy base against changes in hydrology through its balance of trapping and retention. 5. As the frequency of floods increased, the spatial distribution of CPOM became increasingly patchy as storage was driven entirely by structures with high retention. Thus, the coupling of flood frequency and geomorphic structure influenced CPOM availability, which in turn has ramifications for the entire stream food web.     

A novel method reveals how channel retentiveness and stocks of detritus (CPOM) vary among streams differing in bed roughness

1. Coarse particulate organic matter (CPOM) is a fundamental resource in freshwater streams, providing food, shelter and habitat for diverse invertebrate taxa and playing a key role in metabolism in low-order streams. Benthic CPOM stocks are determined by rates of supply and breakdown of detritus and by channel retentiveness (i.e. the capacity for the channel to trap and retain CPOM). We focussed on factors affecting the retentiveness of channels, which theoretically differs among streams with different sediment sizes and concomitant channel morphology.
2. We developed a new, rapid method to measure retentiveness using line-intercept surveys along transects. With this rapid approach, we surveyed 32 sites from three types of streams (smooth sandy channels, n = 10; gravel channels of intermediate roughness, n = 12; rough cobble channels, n = 10) in Victoria, south-eastern Australia, and tested the simple hypotheses that: (1) retentiveness increases in channels with increasing channel roughness (i.e. sandy versus gravel versus cobble-bed streams); (2) different types of channel features (e.g. log jams, cobbles, depositional areas) differ in the efficiency with which they retain CPOM. The line-intercept survey method was readily adapted to measure retentiveness as m of retentive structure per m of transect (i.e. the Linear Coverage Index) and trapping efficiency as m of CPOM per m of retentive element, for 10 different types of retentive elements.
3. Unexpectedly, the retentiveness of channels did not increase with channel roughness. This occurred because channels with different roughness were dominated by different types of retentive structure. Retentive structure in cobble sites was dominated by cobbles themselves, which were highly retentive in other studies but poorly retentive in our system. Gravel and sand sites had more log jams and depositional areas, such as pools and backwaters, and these features were more effective at trapping CPOM. Thus, retention of CPOM was highest in gravel and sand sites.
4. Our method provides a new tool for investigators testing hypotheses about CPOM retention in streams. The method is rapid, requires a minimum of equipment and personnel, and may be applied in any wadeable stream. Retentiveness is calculated in intuitive units that are directly comparable among sites and may have utility as variables in models of CPOM dynamics. We hope this method will open up new avenues for research that may shed light on how CPOM stocks vary among streams, with implications for diversity of aquatic fauna and ecosystem functions such as decomposition.

     

The Effect of Riffle-Scale Environmental Variability on Macroinvertebrate Assemblages in a Tropical Stream

In a tropical stream (at the Soberaní;a National Park, Panama), different environmental factors were quantified in riffle habitats (water characteristics: velocity, depth, turbulence, and direction; stone characteristics: surface area, sphericity, and degree of burial; and others: substrate type, and canopy cover). Characteristics of macroinvertebrate assemblages (mean density of individuals, mean taxon richness, and cumulative taxon richness in three stones at each riffle) were related to both mean values and variability of these environmental factors at riffle scale. Macroinvertebrate density was higher in shallow, fast flowing, stony riffles, with low variability in dominant substrate type. Taxon richness was also higher in shallow riffles with loose, not buried stones, and water direction more or less parallel to the bank. Environmental variability resulted as important as mean values of environmental factors to explain variation in macroinvertebrate assemblages. This is the first study, to our knowledge, that quantifies substratum variability and demonstrates its influence on macroinvertebrate assemblages in a tropical stream.     

Litter in a first–order stream of a temperate deciduous forest (Margaraça Forest, central Portugal)

To evaluate the importance and fate of organic matter inputs in forested streams, we determined the litterfall inputs and the benthic coarse particulate organic matter (CPOM) in one headwater stream flowing through a mixed deciduous forest, during one year. Both vertical traps and the stream bottom were sampled monthly. The material collected was sorted into four main categories: leaves, fruits and flowers, twigs and debris. Litter production was 715 g m−2 y−1 and seasonal, with 73% of the annual total during October–December (autumn). Leaves comprised the largest litter component. Benthic organic matter was 1880 g m−2 y−1, and was also seasonal. Highest accumulation was attained in spring, and twigs and branches comprised the major component. This revised version was published online in July 2006 with corrections to the Cover Date.     

Trophic roles of tadpoles in tropical Australian streams

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Woody Debris Accumulations — Important Ecological Components in a Low Order Forest Stream (Weidlingbach,Lower Austria)

From May 1996 to February 1998, the density of debris dams was surveyed in monthly intervals along a 750-m-long, third order section of the Weidlingbach, a fourth order sandstone brook in the Wienerwald. The mean density over the whole study period was one woody debris accumulation per 35 m stream length. The longevity of these structures ranged from one month to over one year, depending on the hydrology of the stream. More than 50% of the accumulations outlasted moderate floods (0.3–0.4 m³ s⁻¹); of these, 52% were tightly fixed to the bank vegetation. However, only 12% of the destroyed dams were anchored in any way. In addition, two dams were sampled for macroinvertebrate colonisation every other month from June 1996 to June 1997. Dry weights of the leaf and wood fraction were determined and macroinvertebrates identified. Within dams, organismic densities were lowest in temporarily dry levels at the top of debris accumulations (mean density =171 specimens dm⁻³) and highest in permanently submerged levels near the water surface as well as in patches with high leaf litter concentrations (mean density = 1240 specimens dm⁻³). Within the topmost 5 cm sediment layer mean macroinvertebrate density was 67.7 specimens dm⁻³. Within debris dams, taxa composition was not significantly different between the lowermost level 1 and the sediment surface and among levels within dams; however, significant differences existed between the topmost level 4 and all other dam levels. In level 4, we observed significantly more terrestrial Diptera taxa and Coleoptera and significantly fewer Plecoptera and Gammaridae.