Acidic episodes retard the biological recovery of upland British streams from chronic acidification

We tested two predictions required to support the hypothesis that anthropogenic acidic episodes might explain the poor biological response of upland British streams otherwise recovering from acidification: (i) that invertebrate assemblages should differ between episodic and well-buffered streams and (ii) these effects should differentiate between sites with episodes caused by anthropogenic acidification as opposed to base-cation dilution or sea-salt deposition. Chronic and episodically acidic streams were widespread, and episodes reflected acid titration more than dilution. Nonmarine sulphate (16–18% vs. 5–9%), and nitrate (4–6% vs. 1–2%) contributed more to anion loading during episodes in Wales than Scotland, and Welsh streams also had a larger proportion of total stream sulphate from nonmarine sources (64–66% vs. 35–46%). Sea-salts were rarely a major cause of episodic ANC or pH reduction during the events sampled. By contrast, streams with episodes driven by strong anthropogenic acids had lower pH (5.0±0.6) and more dissolved aluminium (288±271 μg L⁻¹) during events than where episodes were caused by dilution (pH 5.4±0.6; 116±110 μg Al L⁻¹) or where streams remained circumneutral (pH 6.7±1.0; 50±45 μg Al L⁻¹). Both biological predictions were supported: invertebrate assemblages differed among sites with different episode chemistry while several acid-sensitive species were absent only where episodes reflected anthropogenic acidification. We conclude that strong acid anions – dominantly nonmarine sulphate – still cause significant episodic acidification in acid-sensitive areas of Britain and may be a sufficient explanation for slow biological recovery in many locations.     

Recovery from acidification in boreal lakes inferred from macroinvertebrates and subfossil chironomids

Acidification of waters and soils caused by emissions and the long-range transport of air pollutants has been a serious worldwide problem during the last decades. The extent of the acidification problem in Finnish acid-sensitive forest lakes was examined in the Acidification Research Project (HAPRO) in the mid-1980s. The recent decline in the emissions of air pollutants has resulted in the chemical recovery of watersheds in many regions, and the present work on the recovery processes in acidified Finnish headwater lakes (REPRO) was launched to examine whether the chemical recovery has already been accompanied by biological recovery. The patterns of recovery were studied by re-sampling littoral macrozoobenthos in a subset of the previously sampled HAPRO lakes. Paleolimnological samples were taken in order to assess the possible dependence of lacustrine chironomid communities on the changing degree of acidification. Acid sensitive and moderately acid sensitive benthic species revealed slight recovery in the formerly most acidic (pH 5.5) but recently recovered lakes. The most significant factors affecting the response of benthic communities were increased mean lake pH and decreased labile aluminium concentration. Paleolimnological chironomid analysis revealed a slight response along the pH gradient, but also significant structural similarity between the present and pristine chironomid assemblages. This implies that no major changes in chironomid communities of these acidic lakes have occurred during the past centuries. The alternative future trends and threats to biological recovery in small headwater lakes are discussed.     

Bioindication in Norwegian rivers using non-diatomaceous benthic algae: The acidification index periphyton (AIP)

Impacts of acidification on aquatic communities have been obvious in Scandinavia since the end of the 19th century and recent model simulations show that in Norway, acidification of surface waters will continue to be an issue in the coming decades. Here, we present a new index based on non-diatomaceous benthic algae (acidification index periphyton, AIP) that can be used to describe the mean annual acidity of Norwegian rivers. The AIP was applied to 608 samples from unlimed rivers all over Norway and values ranged from 5.35 to 7.28, thus covering a range from acid to neutral conditions. Application of the AIP to both limed sites and sites that formerly were acidified demonstrate that the algal community reacts with a several years delay to both river liming and natural recovery. The AIP is most sensitive between mean annual pH values of approximately 5.5 and 7.0 and can be especially useful in detecting the first signs of an acidification trend or the last steps of a recovery process.     

Current velocity influences the facilitation and removal of algae by stream grazers

The modification of flows in lotic ecosystems can have dramatic effects on abiotic and biotic processes and change the structure of basal trophic levels. In high-gradient streams, most of the biota are benthic, and decreased flow may homogenize and reduce benthic current velocity, potentially changing stream ecosystem function. Grazing by macroinvertebrates is an important component of stream function because grazers regulate energy flow from primary producers to higher trophic levels. We conducted an experiment to examine how macroinvertebrate grazers facilitated or removed algal biomass across a gradient of benthic current velocity (0–40 cm s^?1). We chose three grazers (Drunella coloradensis, Cinygmula spp., and Epeorus deceptivus) from a montane stream and conducted our experiment using 24 artificial stream channels that had three treatments: no grazers (control), single-grazer, and combined-grazer treatments. In the absence of grazers, algal biomass increased with benthic current velocity. Grazer treatments differed from the control in that more algal biomass was removed at higher velocities, whereas algal accrual was largely facilitated at low velocities. The transition from facilitation to removal ranged from 4.5 to 5.9 cm s^?1 for individual grazer treatments and occurred at 11.7 cm s^?1 for the combined-grazer treatment. Our data suggest that velocity plays a significant role in the facilitation and removal of algae by macroinvertebrate grazers. Additionally, the patterns revealed here could have general implications for algal accrual in systems where flow is reduced.     

The importance of acid episodes in determining faunal distributions in Welsh streams

SUMMARY. 1. In regional studies of surface-water acidification, annual means of chemical variables are often used to describe differences and change. Outputs from hydrochemical models are often in the form of mean values, which are used in biological models, and these, in turn, are usually derived from responses to mean conditions. Thus, biological forecasts are constrained to ignore the possible effects of the short-term variations in water chemistry which characterize acid streams. This approach requires appraisal.
2. Here, regional Welsh data and daily records from a smaller number of streams were used to investigate the estimation of pH parameters. Variations in aluminium concentration in relation to pH were also assessed. Empirical relationships between invertebrate assemblages, fish populations, mean stream chemistry and measures of fluctuation in pH and aluminium concentration were explored.
3. In general, pH or Al variability and mean pH or Al in Welsh streams were closely related, so that the biological influences of episodes could not easily be separated from those of chronic conditions. Mean pH and mean aluminium concentration were the most effective pH and aluminium statistics used in multivariate models of trout density, which were not improved by including other pH or aluminium variables. For models of invertebrate assemblages based on mean pH or mean aluminium, the inclusion of variables related to episodicity (e.g. pH minimum, aluminium maximum) gave moderate increases in precision.
4. This analysis indicates that it is reasonable to use means of stream chemical variables in biological models of acidification. Consideration of chemical variability could give improvements in some cases, but at the expense of increased model complexity and effort in parameter selection. Nevertheless, we emphasize the need for accurate calibration of both biological and hydrochemical models.     

Differential modification of seawater carbonate chemistry by major coral reef benthic communities

Ocean acidification (OA) resulting from uptake of anthropogenic CO₂ may negatively affect coral reefs by causing decreased rates of biogenic calcification and increased rates of CaCO₃ dissolution and bioerosion. However, in addition to the gradual decrease in seawater pH and Ω _a resulting from anthropogenic activities, seawater carbonate chemistry in these coastal ecosystems is also strongly influenced by the benthic metabolism which can either exacerbate or alleviate OA through net community calcification (NCC = calcification – CaCO₃ dissolution) and net community organic carbon production (NCP = primary production ? respiration). Therefore, to project OA on coral reefs, it is necessary to understand how different benthic communities modify the reef seawater carbonate chemistry. In this study, we used flow-through mesocosms to investigate the modification of seawater carbonate chemistry by benthic metabolism of five distinct reef communities [carbonate sand, crustose coralline algae (CCA), corals, fleshy algae, and a mixed community] under ambient and acidified conditions during summer and winter. The results showed that different communities had distinct influences on carbonate chemistry related to the relative importance of NCC and NCP. Sand, CCA, and corals exerted relatively small influences on seawater pH and Ω _a over diel cycles due to closely balanced NCC and NCP rates, whereas fleshy algae and mixed communities strongly elevated daytime pH and Ω _a due to high NCP rates. Interestingly, the influence on seawater pH at night was relatively small and quite similar across communities. NCC and NCP rates were not significantly affected by short-term acidification, but larger diel variability in pH was observed due to decreased seawater buffering capacity. Except for corals, increased net dissolution was observed at night for all communities under OA, partially buffering against nighttime acidification. Thus, algal-dominated areas of coral reefs and increased net CaCO₃ dissolution may partially counteract reductions in seawater pH associated with anthropogenic OA at the local scale.     

Mechanisms of benthic algal recovery following spates: comparison of simulated and natural events

We conducted a manipulative field experiment to examine individual and interactive effects of scour and short-term nutrient enrichment (4 h exposure) on postspate recovery of benthic algae in a desert stream. We then compared recovery from these simulated-spate conditions to algal recovery patterns following a natural spate that increased water-column nutrient levels for 2 weeks. That event differentially scoured communities on artificial substrata in place for a long-term experiment, significantly reducing biomass in 49-day-old communities but causing no significant reduction of biomass in older, 133-day-old communities. Thus, we were able to examine recovery of scoured and non-scoured benthic algal communities under natural post-spate conditions. Both natural and simulated spates reduced actual and relative abundances of diatoms within communities. In the manipulative experiment, scoured communities accrued biomass more rapidly than those not subjected to scour, but short-term enrichment had not effect. Accrual of diatoms and green algae was stimulated by the scour manipulations, while cyanobacteria maintained equal rates of growth in all treatments. Following the natural spate, diatom and green-algal densities increased in scoured communities, but recovery of algal biomass was slow on both scoured and non-scoured substrata, primarily because cyanobacteria, the dominant algal group on all tiles, did not increase under exposure to highly nitrate-enriched waters. Rates of algal cell accrual were inversely correlated with the amount of algal biomass present at the start of a recovery sequence. Algal immigration rates measured immediately after the natural spate and during an interflood period in the same season did not differ, indicating that the algal drift pool was not augmented by disturbance. Benthic algal recovery following spates is strongly influenced by the degree of scour generated by the event, but recovery patterns are also affected by the length of post-spate enrichment and the taxonomic composition of the affected community.     

NUTRIENT LIMITATION OF BENTHIC ALGAE IN LAKE MICHIGAN: THE ROLE OF SILICA1

In the Laurentian Great Lakes, phytoplankton growth and biomass are secondarily limited by silica (Si), as a result of phosphorus (P) enrichment. Even modest levels of P enrichment can induce secondary Silimitation, which, in turn, promotes a shift from the native diatom phytoplankton flora to chlorophyte and cyanobacteria species. However, very little is known about the nutritional status of benthic populations and their response to nutrient enrichment. Two experiments were performed in the littoral zone of Lake Michigan where nutrients were delivered to in situ benthic algal (episammic and epilithic) assemblages using nutrient‐diffusing substrata. In order to test the hypothesis that benthic algae in Lake Michigan are Si limited, a 2 × 3 factorial experiment was used to deliver all combinations of Si, N, and P to resident assemblages growing on artificial substrata composed of natural (Si rich) versus calcium carbonate (Si poor) sand. A second experiment utilized a serial enrichment to evaluate the role of Si in mediating changes in taxonomic composition. These findings indicate that benthic algae in Lake Michigan exhibit signs of secondary Si limitation, and that their response to enrichment is similar to the phytoplankton. Moreover, natural sand substrata may provide a source of Si to resident benthic algae.     

Benthic algal biomass in an unshaded first-order lowland stream: distribution and regulation

The vertical distribution of algal biomass in the bed sediment and the seasonal development of benthic algae on stones and fine-grained sediments were studied in a small unshaded stream. In addition, field experiments were conducted on the role of irradiance and phosphorus in regulating algal biomass. We found that algal biomass was high at a sediment depth of ten centimetres. Comparison of studies on algal biomass where different depths of the sediment are used should therefore be made with caution. Substrata-dependent differences in algal biomass development were substantial. While algal biomass development on stones was controlled by macroinvertebrate grazing, that on the fine-grained sediment followed the dynamics of incident irradiance, but was attenuated by sediment rebedding. Because of the high grazing pressure on algal biomass on stony substrata, no significant response to phosphorus enrichment was attained. In contrast, algal biomass development on fine-grained sediments was phosphorus-limited. Heavy shading of the fine-grained sediments did not significantly affect algal biomass development, thus suggesting that phosphorus limitation prevents algae from fully utilizing the light resource in this stream. This revised version was published online in July 2006 with corrections to the Cover Date.     

Comparing Effects of Nutrients on Algal Biomass in Streams in Two Regions with Different Disturbance Regimes and with Applications for Developing Nutrient Criteria

Responses of stream algal biomass to nutrient enrichment were studied in two regions where differences in hydrologic variability cause great differences in herbivory. Around northwestern Kentucky (KY) hydrologic variability constrains invertebrate biomass and their effects on algae, but hydrologic stability in Michigan (MI) streams permits accrual of high herbivore densities and herbivory of benthic algae. Multiple indicators of algal biomass and nutrient availability were measured in 104 streams with repeated sampling at each site over a 2−month period. Many measures of algal biomass and nutrient availability were positively correlated in both regions, however the amount of variation explained varied with measures of biomass and nutrient concentration and with region. Indicators of diatom biomass were higher in KY than MI, but were not related to nutrient concentrations in either region. Chl a and % area of substratum covered by Cladophora were positively correlated to nutrient concentrations in both regions. Cladophora responded significantly more to nutrients in MI than KY. Total phosphorus (TP) and total nitrogen (TN) explained similar amounts of variation in algal biomass, and not significantly more variation in biomass than dissolved nutrient concentrations. Low N:P ratios in the benthic algae indicated N as well as P may be limiting their accrual. Most observed responses in benthic algal biomass occurred in nutrient concentrations between 10 and 30 μg TP l⁻¹ and between 400 and 1000 μg TN l⁻¹.     

Impacts of acidification on macroinvertebrate communities in streams of the western Adirondack Mountains,New York,USA

Limited stream chemistry and macroinvertebrate data indicate that acidic deposition has adversely affected benthic macroinvertebrate assemblages in numerous headwater streams of the western Adirondack Mountains of New York. No studies, however, have quantified the effects that acidic deposition and acidification may have had on resident fish and macroinvertebrate communities in streams of the region. As part of the Western Adirondack Stream Survey, water chemistry from 200 streams was sampled five times and macroinvertebrate communities were surveyed once from a subset of 36 streams in the Oswegatchie and Black River Basins during 2003–2005 and evaluated to: (a) document the effects that chronic and episodic acidification have on macroinvertebrate communities across the region, (b) define the relations between acidification and the health of affected species assemblages, and (c) assess indicators and thresholds of biological effects. Concentrations of inorganic Al in 66% of the 200 streams periodically reached concentrations toxic to acid-tolerant biota. A new acid biological assessment profile (acidBAP) index for macroinvertebrates, derived from percent mayfly richness and percent acid-tolerant taxa, was strongly correlated (R² values range from 0.58 to 0.76) with concentrations of inorganic Al, pH, ANC, and base cation surplus (BCS). The BCS and acidBAP index helped remove confounding influences of natural organic acidity and to redefine acidification-effect thresholds and biological-impact categories. AcidBAP scores indicated that macroinvertebrate communities were moderately or severely impacted by acidification in 44–56% of 36 study streams, however, additional data from randomly selected streams is needed to accurately estimate the true percentage of streams in which macroinvertebrate communities are adversely affected in this, or other, regions. As biologically relevant measures of impacts caused by acidification, both BCS and acidBAP may be useful indicators of ecosystem effects and potential recovery at the local and regional scale.     

Influence of the Thin Drift Algal Mats on the Distribution of Macrozoobenthos in Kõiguste Bay, NE Baltic Sea

Macrozoobenthic communities within and outside of the drift algal mats were compared in Kõiguste Bay, NE Baltic Sea. The patches of the drift algae were on average 0.5–1 km wide in diameter covering about 25% of the total bottom area of the bay. Thickness of the mat did not exceed 6 cm. The biomass of the mat varied between 35 and 1391 g dw m^?2. The drift algal mats had no clear negative effect on macrozoobenthos except for a few infaunal species. The drift algae favoured several detrivorous, herbivorous and carnivorous species. Among the studied variables, the thickness of algal mat and oxygen concentration at near-bottom layer explained the best the structure of macrozoobenthos. Total number of invertebrate species increased curvilinearly with the thickness of algal mat having the peak value at 3–5 cm thick algal mat. To conclude, moderate drift algal mats increased habitat complexity and, thus, the diversity of benthic faunal assemblages in otherwise poorly vegetated coastal areas.     

Biomanipulation-induced reduction of sediment phosphorus release in a tropical shallow lake

Biomanipulation via fish regulation combined with submerged plant introduction is an effective measure to restore eutrophic shallow lakes. Improved water quality and clarity promote growth of benthic algae, which with submerged plants may limit sediment phosphorus (P) release, thereby reinforce lake recovery. Our study sought to evaluate the effect of such a biomanipulation on water quality, benthic algal development and sediment P release in a shallow, tropical lake by (1) comparing porewater and lake water quality, light intensity and benthic algal development in restored and unrestored sections; (2) conducting a ³²P radiotracer experiment to track P release from sediment cores sampled from both sections. The biomanipulation led to lower total P, total dissolved P, and soluble reactive P concentrations in lake water, lower phytoplankton biomass, and increased light intensity at sediment surface, stimulating benthic algal development. Moreover, sediment ³²P release was lower in the restored than unrestored section. Concurrently, dissolved oxygen levels in upper layers of the sediment cores were higher in the restored section. Our study indicates that the biomanipulation improved water quality and enhanced growth of benthic algae, thereby reducing sediment P release, which may be one of the main mechanisms to create successful restoration.     

Response of an algal assemblage to nutrient enrichment and shading in a Hawaiian stream

To investigate the effects of nitrate enrichment, phosphate enrichment, and light availability on benthic algae, nutrient-diffusing clay flowerpots were colonized with algae at two sites in a Hawaiian stream during spring and autumn 2002 using a randomized factorial design. The algal assemblage that developed under the experimental conditions was investigated by determining biomass (ash-free dry mass and chlorophyll a concentrations) and composition of the diatom assemblage. In situ pulse amplitude-modulated fluorometry was also used to model photosynthetic rate of the algal assemblage. Algal biomass and maximum photosynthetic rate were significantly higher at the unshaded site than at the shaded site. These parameters were higher at the unshaded site with either nitrate, or to a lesser degree, nitrate plus phosphate enrichment. Analysis of similarity of diatom assemblages showed significant differences between shaded and unshaded sites, as well as between spring and autumn experiments, but not between nutrient treatments. However, several individual species of diatoms responded significantly to nitrate enrichment. These results demonstrate that light availability (shaded vs. unshaded) is the primary limiting factor to algal growth in this stream, with nitrogen as a secondary limiting factor.     

How protective are refuges? Quantifying algal protection in rock crevices

1. Refuges can be functionally important if they harbour sufficient organisms during disturbance to augment population recovery. I quantified the protection of stream algae in crevice refuges using the applied, severe disturbance of scrubbing. Scrubbing effectively removed visible surface algae, and algae remaining on stones were considered protected. 2. In a field experiment, substrata with different quantities of crevice (glass bottles; greywacke; schist; pumice) were incubated in a channelised stream. The possible influence of growth conditions was investigated concurrently; half of the substrata were suspended in the water column, the rest were placed on the bottom (providing differential access to grazers and exposing them to different flow conditions). 3. Rougher substrata had greater total algal biomass than smoother substrata; this pattern resulted from more algal biomass in crevices of the rougher substrata. Protection from scrubbing ranged from about 5% of total algal biomass on glass and greywacke to 80% on pumice. In contrast, algal biomass removed by scrubbing was similar among the experimental substrata. Suspended substrata had more chironomid grazers than those on the bottom, and also greater algal biomass, possibly because of high algal concentrations in the chironomid retreats. 4. A survey of stones from three rivers supported the experimental results; namely, rough pumice protected more algae from scrubbing than did smoother greywacke. 5. In a separate experimental assay, there was no difference in algal growth on agar plates with and without added powdered rock substrata, suggesting that crevice characteristics and not substratum chemistry produced the differences in algal assemblages between rough and smooth surfaces. 6. Results indicate that rough stream stones may protect sufficient algae to augment their recovery in streams following disturbance.     

Assessing the short-term response of stream diatoms to acidity using inter-basin transplantations and chemical diffusing substrates

1. Acid‐base status has major effects on diatoms, but there is little information on their short‐term response to changing acidity. This is despite the use of diatoms as bioindicators in streams where acid episodes are important during rainstorms (hours to days) or snowmelt (days to weeks). In the Llyn Brianne experimental catchments (Wales, UK), we attempted to mimic the effects of short‐term acidification by (i) reciprocally transplanting diatoms between two streams of contrasting acidity and (ii) using acid‐diffusing substrates. 2. Diatom diversity decreased rapidly on substrata transplanted from the circumneutral into the acidic stream, and increased in the reciprocal transplantation. Changes in dominant taxa occurred within three days in the acidic stream because of the rapid growth of Eunotia exigua, and by nine days in the circumneutral stream because of the proliferation of Achnanthidium minutissimum. Transplants were near indistinguishable from ambient assemblages by day 12. 3. There were no effects of enclosures on assemblage composition, but diatoms responded more rapidly to altered chemistry in enclosures with coarse mesh (26 × 50 mm) than finer mesh (320 μm). 4. Chemical diffusing substrates comprised terracotta tiles attached to dosing reservoirs that created locally acid (using H₂SO₄) or metal‐rich conditions (using MnSO₄) in the circumneutral stream over 26 days. Diatom responses were compared with reference substrates dosed with deionised or circumneutral stream water, and we also assessed whether effects were moderated by macroinvertebrate grazers. 5. Surface pH was lower by 1–2 pH units on acid‐dosed substrates than on reference tiles or in surrounding streamwater. Grazed assemblages on acid‐dosed substrates differed significantly from ungrazed reference assemblages, acquiring significantly greater relative abundance of Eunotia spp. However, the magnitude of response was less than in the between‐stream transplantations either because (i) metal exposure and base cation concentrations differed between the transplants and dosing substrates or (ii) diatom response to reduced pH on the diffusing substrates was restricted by the scarcity of acidobiontic diatoms in the circumneutral stream. Similar filter, founder or dominance effects might also affect diatom responses to real acid episodes. 6. These data show that diatom assemblages can respond rapidly and directly to changes in acid‐base status, but short‐term acidification might affect diatoms more rapidly than subsequent recovery. Because the experimental methods used were imperfect representations of episodic effects, diatom response to real acid events requires further field evaluation.     

Exclusion size and material have minimal effects on stream benthic algae and macroinvertebrate colonization within submerged cages

Despite their widespread use in grazer–biofilm studies, stream exclusion cages have inherent physical properties that may alter benthic organism colonization and growth. We used laboratory studies and a field experiment to determine how exclusion cage design (size and material) alters light availability, water velocity, and benthic organism colonization. We measured light reduction by various plastic cage materials and flow boundary layer thickness across a range of exclusion cage sizes in the laboratory. We also deployed multiple exclusion cage designs based on commonly available materials into a second-order stream to assess algae and macroinvertebrate colonization differences among exclusion cages. All plastics reduced some light (190–700 nm wavelengths) and blocked more ultraviolet light than photosynthetically active radiation. Exclusion cage size did not influence flow boundary layer thickness, but larger exclusions tended to have higher velocity at the substrata surface. Despite light and water velocity differences, algal biomass, macroinvertebrate density, and community composition were similar between exclusion cage types. However, algal assemblages outside exclusion cages differed in composition and had higher biomass compared to inside exclusion cages. In terms of algal and macroinvertebrate colonization, plastic exclusion cage size and material appear to be flexible within the sizes tested, but differences can still exist between exclusion cage communities and those within the stream. Overall, artifacts of screened exclusion cages do not appear to introduce large bias in results of grazer–biofilm studies, but efforts to design exclusion cages that better mimic the natural system should continue.     

Succession of algal communities in headwaters: a comparison of pine afforested and natural grassland streams

We evaluated changes in the succession process in benthic algal communities of an afforested stream by comparing them with those in a grassland stream through analysis of temporal changes in composition, structure and functional traits in a 3-month experiment. We hypothesized that sunlight intensity reduction as a result of streamside implanted tree shading reduces the succession rate and community physiognomic complexity. We selected two streams with different riparian cover (grassland and pine afforestation) for installation of unglazed tiles as artificial substrates. The tiles were collected from each stream after 23, 45 and 73 days of exposure. Afforestation produced changes in algal succession, with lower biomass, lower diversity at the beginning of the experiment, and an increase in the proportion of small, low-profile, tolerant, unicellular and stalked algae, coinciding with the predominance of Achnanthidium. However, the grassland stream contained a higher proportion of intermediate-size algae, as well as a larger proportion of high-profile algae, coinciding with the expected results. In the afforested stream, succession proceeded toward the dominance of low-profile species, that were tolerant of low-light conditions. However, in grassland streams with higher sunlight availability, high-profile algae prevailed without the displacement of tolerant forms. Overall, our results indicate that algal communities in afforested stream remain structurally simpler.     

High assemblage persistence in heterogeneous habitats: an experimental test with stream benthic algae

1. The persistence of biological assemblages is positively affected by spatial heterogeneity. This influence may be indirect, through increased species richness. Another possibility is the increased availability of refuges from disturbances, which would prevent local loss. 2. We conducted a field experiment to test the hypothesis that greater roughness (a form of spatial heterogeneity) on the surface of substrata allows higher persistence of assemblages of stream benthic algae and that this relationship does not depend on species richness. Samples were taken on six occasions from smooth and rough artificial substrata used for algal colonisation. We calculated the persistence of assemblages using two analytical approaches: the mean distance to group centroid and the sum of the Euclidean distances between consecutive sampling occasions, both in a multivariate space. We also subsampled the data to take into account differences in species richness between treatments and thus to evaluate the effect of species richness on persistence. 3. Assemblages on rough substrata were more persistent than assemblages on smooth substrata. The effects detected were not due to the greater species richness on rough substrata, since a higher persistence of the assemblages on rough substrata remained after the subsampling procedures. 4. Our results indicate a strong positive relationship between substratum roughness and the persistence of stream benthic algal assemblages. We suggest that this is due to the presence of physical refuges in heterogeneous habitats.     

The ecological effect of acid conditions and precipitation of hydrous metal oxides in a Rocky Mountain stream

Periphyton and benthic invertebrates assemblages were studied at the confluence of two Rocky Mountain streams, Deer Creek and the Snake River near Montezuma, Colorado. Upstream from the confluence the Snake River is acidic and enriched in dissolved trace metals, while Deer Creek is a typical Rocky Mountain stream. In the Snake River, downstream from the confluence, the pH increases and hydrous metal oxides precipitate and cover the streambed. The algal and benthic invertebrate communities in the upstream reaches of the Snake River and in Deer Creek were very different. A liverwort, Scapania undulata var. undulata, was abundant in the Snake River, and although periphyton were very sparse, there were as many benthic invertebrates as in Deer Creek. Downstream from the confleunce, the precipitation of hydrous metal oxides greatly decreased the abundance of periphyton and benthic invertebrates. This study shows that in streams metal precipitates covering the streambed may have a more deleterious effect on stream communities than high metal-ion activities.