Pathways for algal recolonization in seasonally‐flowing streams

1. In semi‐arid climates, seasonally‐flowing streams provide most of the water required for human use, but knowledge of how water extraction affects ecological processes is limited. Predicted alterations in stream flows associated with the impacts of climate change further emphasize the need to understand these processes. Benthic algae are an important base for stream food webs, but we have little knowledge of how algae survive dry periods or respond to altered flow regimes. 2. We sampled 19 streams within the Grampians National Park, south‐eastern Australia and included four components: a survey of different drought refuges (e.g. permanent pools, dry biofilm on stones and dry leaf packs) and associated algal taxa; a survey of algal regrowth on stones after flows recommenced to determine which refuges contributed to regrowth; reciprocal transplant experiments to determine the relative importance of algal drift and regrowth from dry biofilm in recolonization; direct measurement of algal drift to determine taxonomic composition in relation to benthic assemblage composition. 3. Algae showed little specificity for drought refuges but did depend on them; no species were found that were not present in at least one of the perennial pool, dry biofilm or leaf pack refuges. Perennial pools were most closely correlated with the composition of algal assemblages once flows resumed, but the loss or gain of perennial pools that might arise from stream regulation is unlikely to affect the composition of algal regrowth. However, regulated streams were associated with strong increases in algal density in dry biofilm, including increased densities of Cyanobacteria. 4. A model for algal recolonization in seasonally‐flowing streams identified three pathways for algal recolonization (drift‐dependent, dry biofilm‐dependent and contributions from both), depending on whether streams are diatom‐dominated or dominated by filamentous algae. The model predicted the effects of changes to stream flow regimes on benthic algal recolonization and provides a basis for hypotheses testable in streams elsewhere.     

Responses of benthic macroinvertebrates in small intermittent streams to silvicultural practices

We examined macroinvertebrate communities in small(0.1-1.0 m²) pools of intermittent streams (alwayscontainingsome water but without perennial flow) with small watersheds(2-6 ha) subjected to five types of forest harvest to assesspotential impacts of the different harvest methods. Bufferstrips10 m wide were left on each side of the streams. Each harvesttreatment was coupled with a similar unharvested referencestand.An incomplete block design included three 0.05 m² vacuumsamples from each treatment paired with three from theadjacentreferences. There was a high degree of similarity amongreferencesfor parameters other than taxonomic composition (e.g.macroinvertebrate density, number of species, Shannondiversity,functional groups, etc.). Statistically significantdifferenceswere found between references and treatments and among harvestmethods but the responses varied among response variables(density,Shannon-Weiner diversity, species composition), differentspeciesassemblages (all invertebrates, chironomids,Ephemeroptera-Plecoptera-Trichoptera [EPT], isopods), andfunctional group categories (shredders, collector-gatherers).Wecollected 56 taxa, 7–16 per site, with low communitysimilarity(mean Jaccards=0.18, mean Bray-Curtis percentdissimilarity=81). The most severe harvest treatmentsresultedin the highest diversities of total invertebrates in thesesmallspring pool communities.     

Effects of land cover on sediment regime and fish assemblage structure in four southern Appalachian streams

SUMMARY 1. We examined the relationship between catchment land cover, sediment regime and fish assemblage structure in four small streams in the upper Little Tennessee River basin of North Carolina. Study streams drained similar sized catchments (17–31 km²) with different fractions of non-forested land cover. Non-forested land cover was <3% in two 'reference' streams, whereas it was 13 and 22% in two 'disturbed' streams. Land cover data were compared with sediment transport data (suspended and bedload), benthic habitat data (embeddedness, substratum composition and coverage of fines) and fishes collected in autumn 1997.
2. Suspended sediment concentration was significantly higher in disturbed streams during both baseflow and stormflow. During baseflow disturbed streams nearly always exceeded 10 nephelometric turbidity units (NTU), whereas reference streams never exceeded this threshold. The difference in suspended sediment concentration between reference and disturbed streams was more consistent at baseflow than at stormflow. Therefore, baseflow turbidity may be a useful indicator of potential stream degradation.
3. Disturbed sites had five- to nine-fold more bedload transport than reference sites. Both embeddedness and streambed instability increased with increasing non-forested land cover.
4. Relative abundance of fishes requiring clean cobble/gravel substratum for spawning was lower in disturbed streams, whereas relative abundance of mound-building cyprinids, their nest associates and fishes that excavate nests in soft sediments (centrarchids) was higher. Relative abundance of fishes spawning in benthic crevices and gravel (BC + G) declined as the proportion of non-forested land cover increased. This study supports growing evidence that human-induced sedimentation alters stream fish assemblages.     

Independent and interactive effects of nutrients and grazers on benthic algal community structure

Algal responses to nutrients, grazing by Helicopsyche borealis, and concurrent grazing by Helicopsyche and Baetis tricaudatus were examined in recirculating stream chambers. Alagl communities, dominated by Achnanthes minutissima, Cocconeis placentula, and Synedra ulna, were primarily phosphorus-limited. Algal populations responded after only 6 days of nutrient enrichment. Initially, both the adnate diatom Cocconeis and erect diatom Synedra showed positive response to nutrient enrichment. Accumulation of algal biomass between day 3 and 6 in the P enriched treatment was resulted primarily from the growth of Synedra, an overstory rosette-like diatom colony. Such a shift in dominant growth from adnate to erect diatoms is a general phenomenon in periphyton succession in the absence of disturbance. Algal species showed differential responses to an increase of Helicopsyche densities. The accrual rate of Achnanthes continuously decreased with increasing grazer densities. The accrual rates of both Cocconeis and Synedra declined but reached plateaus between medium and high grazing densities. Baetis effectively and exclusively depressed Synedra and had no significant impact on Cocconeis. After concurrent grazing, algal communities were mainly dominated by Cocconeis (approximately 80% of total algal biovolume). The grazer' s mouth structures, grazing efficiencies, and mobility may account for the differential effects of concurrent grazing on algal communities. Significant interactive effects of P and grazing by Helicopsyche indicated that both nutrient addition and grazing may exert significant impact on algal communities. However, grazing may have a much stronger effect on algae than nutrients. Our results indicate that enhancement of algal biomass by P was dampened by grazing activities and that P had no effect on algal biomass in the presence of grazers.     

Effects of Macrobrachium hainanense predation on benthic community functioning in tropical Asian streams

1. Macrobrachium hainanense is a large predatory palaemonid shrimp, present at high densities in pools of low‐order forested streams in Hong Kong. The present study investigated the impacts of M. hainanense on benthic community structure and functions in pools of two streams: Tai Po Kau Forest Stream and Tai Shing Stream. 2. Repeated whole‐pool experiments involving shrimp density manipulations (removal, control and addition) were conducted in both streams between October 2000 and April 2002, and included a wet (May to September) and two dry (October to April) seasons. The three objectives of the study were to determine the effects of M. hainanense predation on benthic macroinvertebrate abundance and species richness, rates of leaf litter breakdown because of effects on detritivores, and periphyton standing stocks by way of an effect on herbivores. 3. Wet season results showed consistent reductions in benthos densities and species richness following heavy rainfall, irrespective of shrimp manipulation. These results suggested that spate‐induced disturbances might override biotic effects and play a dominant role in structuring benthic communities in stream pools in Hong Kong. 4. No significant, reproducible effects on any of the response variables measured in either stream were found during the dry season. Litter breakdown was reduced in the absence of shrimps during one experiment only, suggesting it might be a type I error. These results signified no effect of shrimp removal on benthic communities, or on the functional processes of litter breakdown, or on periphyton accumulation. The large scale of the experimental units (8–40 m²), refuge availability, and the presence of benthic predatory fishes that cropped excess prey made available by removal of M. hainanense, may have contributed to the lack of any effect, despite the abundance of the predatory shrimps.     

Effect of sediment quality on benthic macroinvertebrate communities in streams in the north of France

The Nord/Pas-de-Calais region located in the North of France has been submitted, for several decades, to important pollution due to intensive industrialization. The present study reports on the effects of the metal and organic content of sediments in different streams on the benthic fauna, using multivariate analyses (PCA, CA, co-inertia analysis). Benthic macroinvertebrate communities present different assemblages along the Canal à Grand Gabarit and the river Aa, both in its natural and in its canalized stretches. These differences are more specifically due to anthropic factors such as the regulation of waterways and urban and industrial contamination. Co-inertia analysis enables to bring to the light the existence of a significant co-structure between environmental and faunistic data. This revised version was published online in September 2006 with corrections to the Cover Date.     

Seasonal changes in an intertidal annual algal assemblage in northern Japan: The role of pre-emption and grazing on algal replacement

The pattern and process of seasonal changes in an intertidal annual algal assemblage were examined at Hiura, northern Japan. Short-term field experiments (<2months duration) were set up to quantify the effects of both grazing and pre-emption on species replacement in the assemblage in three different seasons. An 8-month field experiment was set up to quantify long-term effects, including the indirect effects of both grazing and competitive dominance on the community structure. Results suggested that seasonal change in the algal assemblage resulted from the interaction of abiotic environmental change, competition and grazing. The relative contribution of these factors varied within a short period, presumably as a result of seasonal changes in physical environmental stress, free space availability and grazing pressure. From February to March, when grazer density was low and there was much free space available for algae, the dominant species shifted from foliose green alga Monostroma angicava to filamentous red alga Bangia atropurpurea, because B.atropurpurea grew faster than M.angicava. This species replacement was not influenced strongly by biological interaction but by temporal changes in abiotic environmental conditions. From April to mid May, when there was less free space available for algae in the natural community, the dominant B.atropurpurea decreased with increasing foliose red alga Porphyra yezoensis, because only P.yezoensis was able to invade an area pre-empted by algae. Grazing did not affect this species replacement. After mid May, the two dominant species, P.yezoensis and B.atropurpurea, decreased. Their decline was mainly caused by desiccation stress and was partially affected by grazing.     

Scales of spatiotemporal variation in macroinvertebrate assemblage structure in monsoonal streams: the importance of season

1. We assessed the relative importance of different scales of spatial and temporal variability on benthic macroinvertebrate assemblage structure in six unpolluted streams in monsoonal Hong Kong using ordination and complementary multivariate analyses. The spatial scales were regions, sites (streams) and sections (riffles) within sites. The temporal scales were years (three, including one with unusually high rainfall), seasons (dry versus wet) and days within seasons. 2. Significant differences in assemblage structure were manifested at all temporal scales. Those at the site scale were most obvious, whereas demarcation of assemblage structure at the section (riffle) scale was smaller, and there was no significant regional differentiation in assemblage structure. Seasonal variability in assemblage structure was greater than that among years or days. 3. Inter‐year differences in assemblage structure were recorded at all sites, and were noted among all years at some sites but not at others. They were recorded more frequently during the dry season, although their occurrence (in pair‐wise comparisons between years) appeared to be related to differences in the monsoonal (wet season) rainfall. 4. Seasonal differences in assemblage structure were strongly evident at all sites. Inter‐site differences were more apparent during the dry season when local (site‐scale) influences on assemblages were stronger. By contrast, wet‐season samples were more variable because of spate‐induced disturbance, and inter‐site differentiation was less distinct. 5. Differences among days at all sites were relatively minor, but shifts attributable to repeated spate‐induced disturbance were evident at some sites during the wet season. 6. Differences at the section scale were recorded more frequently during the dry season, when the extent of within‐site variability among sections was higher, reflecting increased patchiness within sections resulting from increased substratum heterogeneity and/or greater intensity of biotic interactions. 7. Seasonal shifts in macroinvertebrate assemblage structure at a variety of scales in Hong Kong streams are likely to be attributable to monsoonal rains affecting the relative intensity of abiotic disturbance and biotic interactions in accordance with the harsh‐benign hypothesis.     

Benthic meiofauna assemblage structure of headwater streams: density and distribution of taxa relative to substrate size

Permanent meiofauna taxa and portions of the population of other invertebrates that are temporarily in the meiofauna size class are often precluded from stream studies and assessments. This study was designed to determine the identity, density, and distribution of major meiofauna taxa relative to substrate size in a set of similar headwater streams. Using a coring technique, meiofauna (80 μm–1 mm) and substrate samples were collected from 11 Ozark headwater streams in the Boston Mountain ecoregion of Arkansas, USA. Mean meiofauna density among streams was 1739 ± 436 organisms per l. Permanent meiofauna taxa (Copepoda, Cladocera, Ostracoda, Rotifera, Nematoda, Hydrachnida, and Tardigrada) comprised 22.5% of the organisms collected with a mean density of 394 ± 233 organisms per l; temporary meiofauna taxa (Oligochaeta, Turbellaria, Hydroidea, Chironomidae, Ephemeroptera, and other insects) comprised the remainder with a density of 1346 ± 308 organisms per l. Chironomidae was the most numerous temporary meiofauna taxon, and Hydrachnida was the most numerous permanent taxon. Streams were found to differ significantly in substrate composition and densities of major taxonomic categories. Substrate size was found to predict densities for most of these taxonomic categories. Meiofauna patchiness was reflected in high variability within streams. Canonical correspondence analysis revealed positive associations between Copepoda and Nematoda and silt, and between Copepoda, Nematoda, and Rotifera and fine sand. Hydrachnida and Rotifera were negatively associated with silt and coarse sand, respectively. The potential value of inclusion of meiofauna in stream environmental assessments is discussed.     

The effects of fire on ant trophic assemblage and sex allocation

Fire plays a key role in ecosystem dynamics worldwide, altering energy flows and species community structure and composition. However, the functional mechanisms underlying these effects are not well understood. Many ground‐dwelling animal species can shelter themselves from exposure to heat and therefore rarely suffer direct mortality. However, fire‐induced alterations to the environment may change a species' relative trophic level within a food web and its mode of foraging. We assessed how fire could affect ant resource utilization at different scales in a Mediterranean forest. First, we conducted isotopic analyses on entire ant species assemblages and their potential food resources, which included plants and other arthropods, in burned and unburned plots 1 year postfire. Second, we measured the production of males and females by nests of a fire‐resilient species, Aphaenogaster gibbosa, and analyzed the differences in isotopic values among workers, males, and females to test whether fire constrained resource allocation. We found that, in spite of major modifications in biotic and abiotic conditions, fire had little impact on the relative trophic position of ant species. The studied assemblage was composed of species with a wide array of diets. They ranged from being mostly herbivorous to completely omnivorous, and a given species' trophic level was the same in burned and unburned plots. In A. gibbosa nests, sexuals had greater δ¹⁵N values than workers in both burned and unburned plots, which suggests that the former had a more protein‐rich diet than the latter. Fire also appeared to have a major effect on A. gibbosa sex allocation: The proportion of nests that produced male brood was greater on burned zones, as was the mean number of males produced per nest with the same reproductive investment . Our results show that generalist ants with relatively broad diets maintained a constant trophic position, even following a major disturbance like fire. However, the dramatically reduced production of females on burned zones compared to unburned zones 1 year postfire may result in considerably reduced recruitment of new colonies in the mid to long term, which could yield genetic bottlenecks and founder effects. Our study paves the way for future functional analyses of fire‐induced modifications in ant populations and communities.     

Seasonal variation in benthic invertebrate recolonization of small-scale disturbances in a subtropical Florida lake

Approximately 11.5% of the littoral zone of a hypereutrophic Florida lake is disturbed by the construction of spawning beds by the cichlid fish, Sarotherodon aurea, during a single spawning season. Simulations of these beds were created during three seasons of the year to test for variation in recolonization rates and patterns in the benthic macroinvertebrate community.The seasonal variation in densities of benthic invertebrate populations suggests a direct relationship between reproductive activity and recolonization rate. Recolonization rates of the numerically dominant species, Polypedilum halterale (Diptera: Chironomidae), were much more rapid in the spring and summer than in the winter. In contrast, Hyalella azteca and Glyptotendipes paripes attained peak population densities during the winter season when Polypedilum was present in low densities. These organisms invaded the disturbed area in unusually high abundances and later declined to background levels. The final result was a winter population with densities comparable to the control (undisturbed) area, but the species composition was different.Similarity between disturbed and control communities during the winter season only reached 67% a month after disturbance, while communities during the warmer months attained nearly 90% similarity in less than 15 days. This lack of similarity during the winter indicates that disturbance, at the appropriate time, may play a role in community organization.     

Pollution and tidal benthic communities of the James River Estuary,Virginia

Distribution of benthic communities in the estuarine portion of the James River was controlled mainly by salinity. Pollution effects were localized and difficult to assess because of a rigorous physical environment. Mesohaline and oligohaline communities were very similar to those in other estuaries of the eastern United States. Macrobenthic densities were most severely depressed in tidal freshwater habitats near Richmond & Hopewell, where the major portion of the pollution load enters the river. Cluster analysis of species distributional patterns and ordination of pollution and physical parameters produced similar results, dividing the river into mesohaline, oligohaline, and upper and lower tidal freshwater zones. Further analysis of only the tidal freshwater portion indicated the distribution of benthic communities reflected the location and concentration of pollution sources along the river. Tidal freshwater communities were dominated by the Asiatic clam, Corbicula fluminea, tubificid oligochaetes of the genus Limnodrilus and the chironomid insect larva Coelotanypus scapularis. The fauna of the freshwater zones was very eurytopic with respect to sediment type and has a great resemblance to the fauna of eutrophic lakes. The classical concept of a sharp increase in number of species occurring from oligohaline to freshwater zones was found to be misleading. This increase does not occur until free flowing (or lotic) freshwater areas of greater habitat diversity are reached.     

Assessing scales of variability in benthic diatom community structure

Sources of variability such as sampling method, sample preparation, and sample analysis (taxonomy) might affect our ability to measure differences in community structure between sites in environmental effects studies. It is therefore important to ensure that changes in community structure related to the physical or chemical differences between sites are not hidden by other sources of variability within a site. The goal of this study was to quantify the amount of variability in benthic diatom community structure related to sampling and laboratory procedures. Our results showed that variability in community structure was minimal among replicate microscope slides (< 1%) and among samples collected within a site (1.8%). Variability in community structure was substantially higher between sites located in one stream (16.6%), and highest across streams (59.6%). This suggests that field sampling and laboratory methods do not contribute a large amount of variation into our analyses of benthic diatom community structure across sites.     

Species richness and assemblage structure of Trichoptera in Danish streams

• 1 Species richness and assemblages of Trichoptera from 157 ‘pristine’ Danish lowland stream sites were analyzed, for dependence on geographical position of the sites and simple physical variables, using two way indicator species analysis and detrended correspondence analysis.
• 2 A total of 106 species were recorded, representing ≈ 90% of the species pool known from Danish streams. Only seven species occurred at more than half the sites, whereas an additional 11 species were found at more than a quarter of the sites.
• 3 Although sites showed significant regional differences in environmental variables (stream order, width, slope and presence/absence of riparian forest), species richness and assemblages were primarily correlated with stream order, width and slope. Maximum richness was found at the largest (5th order) stream sites.
• 4 Regional differences in species assemblages were found, with several species being absent from the islands Funen and Bornholm. Species assemblages also differed between forested and non‐forested stream sites.
• 5 We found evidence that stream temperature may be of only minor importance in determining Trichoptera species richness and assemblage composition in Danish streams compared to other size‐related physical factors.

     

Spatial and seasonal variations in benthic algal assemblages in streams in monsoonal Hong Kong

Samples from stone surfaces were collected in pools within four unpolluted hillstreams (two shaded and two unshaded) in monsoonal Hong Kong (lat. 23°N) to elucidate the extent of spatial (within and among streams) and temporal (seasonal) variations in algal biomass and assemblage composition. Sampling continued for over 12 months, incorporating the dry season when streams were at baseflow, and the wet season when spates were frequent. We anticipated that algal biomass would be lower in shaded streams and during the wet season, with associated seasonal differences in assemblage composition or relative abundance of different growth forms (e.g. erect versus prostrate). Benthic chlorophyll a (a proxy for algal biomass) varied among streams from an annual mean of 11.0–22.3 mg m⁻². Dry-season standing stocks were 18% higher than during the wet season when spate-induced disturbance reduced algal standing stocks. Algal biomass varied significantly at the stream scale, but not at the pool scale, and was lower in unshaded streams, where standing stocks may have been limited by high densities of algivorous balitorid loaches (mainly Pseudogastromyzon myersi). An overriding effect of grazers on algal biomass could also have reduced variations resulting from spate-induced disturbance. Significant differences in assemblage composition among streams, which were dominated by diatoms and cyanobacteria (totally 82 taxa) were not systematically related to shading conditions. Seasonal variations in algal assemblages were statistically significant but rather minor, and did not involve major shifts in composition or growth form caused by spate-induced disturbance. The abundance of filamentous cyanobacteria in all the streams may have been due to ‘gardening’ by balitorid loaches that removed erect or stalked diatoms and favoured cyanobacteria that persist through basal regeneration of filaments. This explanation requires validation through manipulative experiments. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users. Handling editor: Luis Mauricio Bini     

Responses of benthic algal communities and their traits to experimental changes in fine sediments,nutrients and flow

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Effect of periphyton biomass on hydraulic characteristics and nutrient cycling in streams

The effect of periphyton biomass on hydraulic characteristics and nutrient cycling was studied in laboratory streams with and without snail herbivores. Hydraulic characteristics, such as average water velocity, dispersion coefficients, and relative volume of transient storage zones (zones of stationary water), were quantified by performing short-term injections of a conservative tracer and fitting an advection-dispersion model to the conservative tracer concentration profile downstream from the injection site. Nutrient cycling was quantified by measuring two indices: (1) uptake rate of phosphorus from stream water normalized to gross primary production (GPP), a surrogate measure of total P demand, and (2) turnover rate of phosphorus in the periphyton matrix. These measures indicate the importance of internal cycling (within the periphyton matrix) in meeting the P demands of periphyton. Dense growths of filamentous diatoms and blue-green algae accumulated in the streams with no snails (high-biomass streams), whereas the periphyton communities in streams with snails consisted almost entirely of a thin layer of basal cells of Stigeoclonium sp. (low-biomass streams). Dispersion coefficients were significantly greater and transient storage zones were significantly larger in the high-biomass streams compared to the low-biomass streams. Rates of GPP-normalized P uptake from water and rates of P turnover in periphyton were significantly lower in high biomass than in low biomass periphyton communities, suggesting that a greater fraction of the P demand was met by recycling in the high biomass communities. Increases in streamwater P concentration significantly increased GPP-normalized P uptake in high biomass communities, suggesting diffusion limitation of nutrient transfer from stream water to algal cells in these communities. Our results demonstrate that accumulations of periphyton biomass can alter the hydraulic characteristics of streams, particularly by increasing transient storage zones, and can increase internal nutrient cycling. They suggest a close coupling of hydraulic characteristics and nutrient cycling processes in stream ecosystems.