Long-term effects of PO4 fertilization on the distribution of bryophytes in an arctic river

1. Each year since 1983, H₃PO₄ has been added continuously during the ice-free season to a P-limited tundra river (Kuparuk River, North Slope, Alaska). Effects on epilithic metabolism, invertebrate community structure and fish production developed quickly. 2. In 1990, 7 years after fertilization began, we noted extensive coverage by bryophytes within the fertilized reach of the river, where very little had been noted before. Bryophyte biomass from a limited set of quadrats taken in 1990 and 1991 yielded 17 ± 9 (SE) g dry mass m^?2 in control reaches and 322 ± 96 g dry mass m^?2 in fertilized reaches. 3. An initial survey of macroalgal and bryophyte cover in 1991 suggested that the moss Schistidium (Grimmia) agassizii was distributed in both control and fertilized reaches of the river. No clear difference in coverage by this species was found in either reach. 4. In contrast, two species of Hygrohypnum (H. alpestre and H. ochraceum) were found almost exclusively in the fertilized reach. An extensive point transect survey done in 1992, above, within and below the fertilized reach, indicated that increased cover and biomass of Hygrohypnum spp. were confined to the fertilized reach of the river. Detrended correspondence analysis clearly separated the macrophyte and macroalgal communities in the fertilized reach from those in the control and downstream reaches. 5. A fourth bryophyte species (Fontinalis neomexicana) also occurred almost exclusively in the fertilized reach, but was much less abundant than the Hygrohypnum species. 6. Analysis of total N and P in the tissues of the Hygrohypnum spp., and estimates of average coverage (～15%) and biomass (～150g dry weight m^?2) over an 8km fertilized reach, suggest that these species alone may have removed two-thirds of the P added in the fertilizer experiment. The bryophyte community in this river is likely to be the dominant sink for P in the fertilized reach.     

Riffles as barriers to interpool movement by three cyprinids (Notropis boops,Campostoma anomalum and Cyprinella venusta)

1. The effects of riffles as barriers to movement of stream fish was investigated in a set of eight large outdoor artificial streams. Pools were 183 cm in diameter and 45 cm deep; riffles were 183 cm long and 43 cm wide. Rates of movement of three species of minnows (Cyprinidae) (Campostoma anomalum, Cyprinella venusta and Notropis boops) among pools were measured at four riffle current velocities (0, 15, 30 and 45 cm s^–1), three thalweg depths (10, 50 and 100 mm), two riffle lengths (183 and 549 cm), and with and without the threat of predation. Visual observation and video cameras were used to quantify movement rate. 2. Mean movement rate (percentage of fish crossing a riffle each 30 min) was 18.1% at 0 cm s^–1 and only 1.8 at 45 cm s^–1. Movement rate was 7.2% with no predators present and 20.2% with caged predators in pools. Notropis boops had a lower rate of movement than C. venusta or C. anomalum across all trials. The mean group size (number of individuals crossing a riffle together) was 1.2 fish overall, indicating most movement was by individuals and not groups. Group size was significantly greater only with shallow riffles or under the threat of predation. 3. Overall biotic and abiotic factors in these artificial streams do influence movement rates and may affect movement among pools in natural streams.     

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.     

Colonization of leaf patches at topographically different locations by insect shredders in a small mountain stream

We examined physical constraints on the colonization of leaf patches by shredder individuals by comparing the colonizations of artificially standardized leaf patches placed at different locations within a stream reach (i.e., riffles, middles and edges of pools). Stonefly taxa (Nemoura, Protonemura) colonized riffle patches 2–10 times more often than pool (middle, edge) patches, whereas caddisfly taxa (two species of Lepidostoma, Nothopsyche) almost exclusively colonized pool patches. Colonization also differed between the middle and edge patches in pools for most taxa; it was 2–5 times greater in edge patches for Nemoura and in middle patches for Lepidostoma. The abilities of species to cope with low oxygen circulation and high shear stress appear to determine differences in colonization between riffle and pool patches, whereas species-specific dispersion behavior (e.g., return time from drift) may differentiate colonization between middle and edge patches in pools. Our results suggest that changes in leaf distribution within a reach can affect the suitability of stream reaches in terms of food acquisition for shredder individuals.     

Crayfish as geomorphic agents and ecosystem engineers: effect of a biomass gradient on baseflow and flood-induced transport of gravel and sand in experimental streams

SUMMARY 1. Using experimental streams, we studied the impact of the crayfish Orconectes limosus on (i) the transport of gravel and sand at baseflow; (ii) the sediment surface (bedform, particle consolidation, proportion of sand, algal and gravel cover); and (iii) the critical shear stress (τ_C) causing incipient gravel and sand motion during simulated floods. We examined (i) and (ii) in experimental outdoor flumes that replicated riffle‐pool sequences and (iii) in a larger laboratory flume, in which we exposed sediments retrieved from the outdoor flumes to a progressively increasing discharge. 2. Habitat changes induced by crayfish, such as bedform alterations in riffles (downstream displacement of riffle heads) and the increase of gravel on sand dunes in pools, had major impacts on the spatial and temporal patterns of the baseflow transport of gravel and sand. 3. In addition to their impact on bedform in riffles and on gravel cover in pools, crayfish prevented the physical consolidation of particles in riffles and reduced the algal cover and the proportion of sand in the surface layer in both riffles and pools. These crayfish impacts on sediment surface variables had complex, interacting effects on the mobilisation of gravel and sand during subsequent flood simulations. For sand, crayfish progressively decreased the τ_C (i.e. the sum of bedform drag and skin friction) by about 50% along the entire biomass gradient in pools, whereas the presence of crayfish abruptly decreased the τ_C by about 75% in riffles. For gravel, the discharge causing motion in riffles produced a shear stress (in terms of skin friction) on an even bedform that was about 75% lower in all flumes with crayfish compared with the flumes without crayfish. Crayfish had no impact on τ_C for gravel in pools. 4. Scaling‐up these experimental results to real streams suggests that crayfish could affect the patch dynamics of major sediment transport events and habitat suitability for other organisms that, at larger spatial scales, could increase the overall spatio‐temporal habitat diversity and thus the overall structural and functional biodiversity of lotic communities.     

Do pools impede drift dispersal by stream insects?

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Reproductive mode of the geographic parthenogenetic mayfly Ephoron shigae, with findings from some new localities (Insecta: Ephemeroptera, Polymitarcyidae)

The burrowing polymitarcyid mayfly Ephoron shigae is distributed widely in Japan. Some populations are bisexual, others are unisexual, and the distributions of the two types overlap broadly. Experimental evidence of parthenogenetic reproduction, long suspected in unisexual populations, is presented here, based on a comparative analysis of the developmental rate of fertilized and unfertilized eggs. The developmental rate of fertilized eggs from 20 mated females in a bisexual population was 98.4% ± 0.73% (mean ± SD), and no unfertilized eggs from 20 virgin females in that population developed. The developmental rates of unfertilized eggs in two unisexual populations were 89.0% ± 4.59% and 84.2% ± 1.96%, respectively. This article presents experimental evidence of geographic parthenogenesis in E. shigae and provides support for the previous interpretation. In addition, we discuss the relationship between the sex ratio of each population and the developmental rate of fertilized versus unfertilized eggs from the females in those populations.     

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。     

The spatial structure of variation in salamander survival,body condition and morphology in a headwater stream network

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Inter-annual changes in the benthic community structure of riffles and pools in reaches of contrasting gradient

The inter-annual variation in the structure of the benthic community of riffles and pools was evaluated in contrasting geomorphic settings. The community structure of riffles and pools was a function of habitat, reach gradient, and discharge and was taxon specific. In years of below average peak discharge, riffles had higher taxon richness than pools (66 versus47) but richness was similar between habitats during a year of average discharge (56 versus 54). The percentage composition of oligochaetes and elmid beetles was more variable inter-annually in pools and low gradient reaches than in high gradient reaches. Differences in the percentage of collector-gatherers and scrapers in riffles and pools appeared related to inter-annual differences in discharge regimes. Two components of the annual discharge regime appear to differentially affect the composition of the benthic community in the snowmelt dominated stream studied: the magnitude of the annual peak discharge and the duration and timing of the period of extended high flow.     

Differences in amounts of pools and riffles between upper and lower reaches of a fully sedimented dam in a mountain gravel-bed river

To understand the extent of changes in the habitat structure of macroinvertebrates due to channel degradation, the amount of pools and riffles and bed materials were determined in upper and lower reaches of a fully sedimented dam in a mountain gravel-bed river. Pools occurred more frequently and individual pools were longer in the lower reach than in the upper reaches; the total length of pools in the former was four times than in the latter. In contrast, individual riffles in the upper reaches were longer than and the total length double that in the lower reach. The main bed materials in the upper reaches were moderate-sized cobbles and boulders (diameter 10–40 cm), which generated moderately steep and long riffles. In the lower reach, these materials were scarce and the main bed materials were large boulders (>50 cm) and bedrock, which generated steep and short riffles; finer materials (<10 cm) dominated in pools and runs. The reduced total length of riffles in the lower reach was partly associated with the steepening of individual riffles, which may be due to the shortage of moderate-sized cobbles and boulders. Macroinvertebrate biomass at reach scale, which was estimated based on the total lengths of pools and riffles, was more than two to four times in the upper reaches than in the lower reach.     

Maternal behaviours may explain riffle‐scale variations in some stream insect populations

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Macroinvertebrate colonization of replacement substrate below a hypolimnial release reservoir

Loss of substrate from occasional high flows below reservoirs on the Trinity River prompted government agencies to replace the gravels to protect the anadromous fisheries resource. Several riffles were restored using 2.5 to to cm gravels. Three of these and two natural riffles, with gravels ranging up to 20 cm, were surveyed over a two-year period to determine the effect of this gravel replacement on the macroinvertebrate fauna. Replaced gravels were too small to support the diversity of organisms found on natural riffle gravels. In addition, temperature alteration by the hypolimnial release reservoir prevented colonization by all but a few organisms able to tolerate such conditions in the riffles below the dam. Large but unstable populations of Simulium, Baetis, and the Chironominae developed below the dam, aided by an abundant periphyton and detrital food supply and lack of competition and predation. It is concluded that habitat diversity through the use of a range of gravel sizes up to the size naturally found in productive riffles should be used in riffle restoration projects in order to increase habitat complexity, species diversity, density, and biomass important in maintaining fishery stocks.     

Spatial distribution and population dynamics of a benthic rotifer, Embata laticeps (Murray) (Rotifera, Bdelloidea) in the bed sediments of a gravel brook

• 1 The spatial and temporal distributions of the most abundant bdelloid rotifer, Embata laticeps (Murray), inhabiting the sediment surface and the hyporheic interstitial of a gravel stream were investigated between October 1991 and October 1992. Three temporal peaks of population density occurred during the year at the sediment surface differing in their riming from the density peaks detected within the bed sediments.
• 2 The depth distributions of E. laticeps in pool and riffle areas differed significantly. Higher abundances were found between 10 and 20 cm depth in the pool and between 20 and 30 cm in the riffle area. In these two sites a significant effect of surface discharge (estimated 1 week before sampling) on rotifer densities was found. Discharge greater than 0.424 m³s^?1 significantly reduced the densities in the riffle, while only at values greater than 1.660m³s^?1 were abundances influenced in the pool area.
• 3 The percentages of egg-bearing females were higher within the bed sediment than in the surface layer, and in the pool area than in the riffle. Thus, overall mean values of population increase were higher in the pool habitat (x?= 0.015 ± SE 0.029 day^?1) than in the riffle and at the sediment surface (x?= 0.004 ± 0.025 and 0.005 ± 0.026 day^?1, respectively). In addition, population growth fluctuated more strongly at the sediment surface than within the hyporheic intersiritial. Comparison of the observed values of population increase with a randomization test, where observed densities were randomly permuted, revealed no significant differences between those observed and values from a random model. Thus, a clear trend in population growth of E. laticeps over time could not be demonstrated in this gravel stream.

     

Ecology and population structure of the bayou darter, <Emphasis Type="Italic">Etheostoma rubrum</Emphasis>: disjunct riffle habitats and downstream transport of larvae

The bayou darter, Etheostoma rubrum (Percidae), is endemic to the Bayou Pierre system in Mississippi. Adult and juvenile E. rubrum occupy swift, shallow riffles or runs over coarse gravel and pebble substrata. Habitat requirements of larval and post-larval stages, and the role of downstream dispersal of larvae in colonizing riffles are poorly known. The potential for movement and the high level of habitat specificity for the discontinuous riffle habitat suggest that E. rubrum may comprise a metapopulation of linearly arranged local populations. The greatest population densities of E. rubrum occur in the upper reaches of Bayou Pierre. We hypothesized that metapopulation structure of E. rubrum may include source–sink dynamics, whereby downstream areas are a sink for larvae/early juveniles originating upstream. We tested hypotheses that a transport mechanism, larval drift, occurred in E. rubrum, and that downstream riffles showed characteristics of population sinks. We captured larval stages of E. rubrum in cross-sectional and longitudinal drift nets, and rates of drift tended to increase during the day. Larval E. rubrum (n=19) occurred in samples above and below riffle areas, with no differences among longitudinal drift nets placed above and below riffles. Thus, larval drift is a viable transport mechanism. Relative abundance of adults and juveniles declined from upstream to downstream, and inter-riffle distances increased with cumulative downstream distance. However, both predictions of the source–sink hypothesis were not supported. The distribution of size classes did not change between upstream and downstream riffles nor did the mean size-adjusted body mass.     

Two lowland stream riffles – linkages between physical habitats and macroinvertebrates across multiple spatial scales

The physical structure of two riffles in a lowland Danish stream was studied and its importance for the composition and density of the macroinvertebrate communities was evaluated. The two riffles were visually assessed to be very similar, but measurements revealed that they differed in overall hydraulic conditions, stability, substratum composition and consolidation. Differences affected abundance of both burrowing and surface dwelling macroinvertebrates. The unstable unconsolidated riffle had higher total macroinvertebrate abundance (4137 m⁻² vs. 1698 m⁻²), diptera abundance (2329 m⁻² vs. 386 m⁻²) and total estimated species richness (31.7 vs. 28.8) as well as lower evenness (0.77 vs. 0.83) than the compact riffle. Among samples within the unconsolidated riffle, variations in macroinvertebrate communities were related to differences in mean substratum particle size. Here a linear log–log relationship existed between macroinvertebrate abundance, the abundance of EPT taxa and the median particle size (r ² _total = 0.46, p = 0.002; r ² _EPT = 0.73, p < 0.001). No similar relationships were evident on the consolidated riffle. Moreover, macroinvertebrate communities on the unconsolidated riffle were dominated by species with a high colonising potential. Despite being assessed to the same morphological unit, physical variation between riffles was surprisingly high as the riffles differed substantially with respect to consolidation, substratum heterogeneity and overall hydraulic structure. Macroinvertebrate community structure and composition also differed between riffles despite being drawn from the same species pool. The findings address the question if we use the correct methods and parameters when assessing the macroinvertebrate communities at the scale of the morphological unit.     

Responses of Fish to Experimental Introduction of Structural Woody Habitat in Riffles and Pools

The declining condition of river systems associated with rapid development of human societies has led to substantial declines in fish diversity. One cause of decline is the loss of in‐stream Structural Woody Habitat (SWH), an important component of stream ecosystems, particularly as fish habitat. As a result there has been an increase in the number of rehabilitation programs that introduce SWH into rivers. This paper assesses fish responses to SWH introduction in riffles and pools in the Hunter River, eastern Australia, using a Multiple‐Before‐After‐Control‐Impact (MBACI) experimental and analytical design. In the riffle experiment, species richness was comparable among all control and treatment riffles across the entire study period. However, there were significant differences in assemblage structure, fish abundance, and biomass between control and treated riffles. The introduction of SWH (bank embedded deflector jams) appeared to create additional habitat which was utilized by one native fish species (Retropinna semoni—Australian smelt) and one alien species (Gambusia holbrooki—mosquito fish). In pools there were no significant changes in fish species richness, abundance, or biomass following introduction of SWH (pool jams). These findings have important practical and cost implications in terms of the design and implementation of rehabilitation strategies using SWH to restore fish assemblages in degraded streams.     

Habitat-specific estimates of competition in stream salmonids: A field test of the isodar model of habitat selection

Summary The population densities of sympatric Atlantic salmon,Salmo salar and brook charr,Salvelinus fontinalis, were measured in riffle and pool stream habitats to test whether non-linear isodars, a multispecific model of habitat selection based on ideal distribution assumptions, could (1) predict the distribution of densities between habitats and (2) reproduce the processes postulated to underlie spatial segregation and species interactions in previous laboratory and field studies. The model provided a good fit to observed density patterns and indicated that habitat suitability declined non-linearly with increased heterospecific competitor densities. Competitive effects in riffles appeared to be due to exploitative resource use, with salmon always emerging as the superior competitor. No evidence was found for interference competition in riffles. In contrast, interspecific competition in pools seemed to occur through exploitation and interference. The specific identity of the superior competitor in pools depended on the density of both species; pools provided the charr with refuge from competition with the salmon, presumably through the adoption by the charr of density-dependent behaviours, such as schooling and group foraging, that mitigated the negative impact of the salmon. Charr were displaced from the riffles toward the pools as the total salmon density increased. The isodar analysis, based on limited density data, successfully reproduced the processes suggested to underlie spatial segregation in previous field and laboratory studies and provided new insights into how changes in competitor densities modify habitat suitability in this system.     

Effect of fertilized,unfertilized, and UV‐irradiated hosts on parasitism and suitability for Trichogramma parasitoids

Trichogramma spp. (Hymenoptera: Trichogrammatidae) parasitoids have been commonly used as biological control agents in insect pest management. Host quality is believed to influence parasitism, host preference, and suitability for parasitoids. To date, limited studies have compared the parasitism of Trichogramma parasitoids on fertilized, unfertilized, and sterilized host eggs. Hence, we studied the performance of three Trichogramma egg parasitoids, Trichogramma japonicum Ashmead, Trichogramma chilonis Ishii, and Trichogramma leucaniae Pang & Chen, on fertilized, unfertilized, and ultraviolet (UV)‐irradiated fertilized (UVF) eggs of rice moth, Corcyra cephalonica (Stainton) (Lepidoptera: Pyralidae). In a no‐choice test, T. japonicum and T. leucaniae parasitized significantly more fertilized or UVF than unfertilized hosts, and T. chilonis parasitized significantly more UVF than either fertilized or unfertilized hosts. In a choice test, all three Trichogramma parasitoids parasitized UVF hosts the most, and unfertilized hosts the least. There were similar percentages of adult emergence and female progeny among fertilized, unfertilized, and UVF hosts for all three Trichogramma parasitoids, except that T. japonicum had significantly lower adult emergence on fertilized hosts. We also found that all three Trichogramma parasitoids developed slower on unfertilized hosts. Regardless of host treatments, T. leucaniae had the longest developmental time and T. chilonis had the shortest. We conclude that Trichogramma parasitoids prefer parasitizing UVF eggs of C. cephalonica without negative effects on their emergence and sex allocation.     

Microfaunal response to fertilization of an arctic tundra stream

1. As part of a whole-system study, the response of the heterotrophic microfaunal community colonizing artificial substrata (polyfoam units) to fertilization of an arctic tundra stream was followed for 6 weeks during the summer. 2. Dominant heterotrophic microfauna observed included amoebae (approximately 40% of colonizing biomass), rotifers (36% of biomass) and ciliates (25% of biomass). 3. Biomass of heterotrophic microfauna on artificial substrata was not significantly different in a control reach and an experimental reach fertilized with phosphorus (loading rate ten times ambient), but in a reach fertilized with both phosphorus and nitrogen (loading rates ten times ambient) biomass was double that of the control and phosphorus-fertilized reaches. The lack of response in the phosphorus reach was probably due to greater insect grazing as a result of previous phosphorus fertilization of this reach. 4. Abundance of microfauna on epilithic surfaces in the river was higher on rocks from pools than on rocks from riffle areas, but abundance on the artificial substrata was higher than on the natural rocks. 5. The results suggest that microfauna of arctic tundra streams are regulated by grazers and that their importance in transfers among trophic levels is greater in pools than in riffles.