Herbivory in an acid stream

• 1 Spatial and temporal variation in the distribution and feeding of non‐predatory macroinvertebrates was investigated in a first‐order, acid stream in the Ashdown Forest, southern England.
• 2 Stonefly (Nemouridae) and chironomid (Orthocladiinae) larvae were abundant on the upper surfaces of mineral substrata of three sizes (small stones, large stones, bedrock). The density of larvae in each taxonomic group did not vary among substrata of different sizes, although strong seasonal variation existed.
• 3 Nemourids and chironomids (H. marcidus) collected from the upper surfaces of substrata exhibited generalist feeding habits, consuming algae (diatoms, coccoid and filamentous green algae), detritus (biofilm matrix material and fine particulate organic matter (FPOM)) and inorganic debris.
• 4 There was spatial variation in the gut contents of nemourids. The proportion of algae in the guts of larvae often increased with the size of the substratum from which they were collected. Strong temporal variation in the composition of the diet also existed. Nemourids ingested a large quantity of attached algae and biofilm matrix from the biofilm in spring and winter, but consumed loose FPOM and associated microflora in summer and autumn.
• 5 We conclude that, in this acid stream, the trophic linkage between algae and grazers is maintained by ‘detritivorous’ stonefly and chironomid species. The relationship between the feeding habits of these larvae and other life‐history attributes, such as mouthpart morphology and mobility, is discussed.

     

Three-year study of benthic algal spring bloom development in a small, Danish lowland stream

Seasonal development of benthic algae was studied over a three-year period in a small, nutrient-rich lowland stream to investigate inter-annual variation in the algal spring bloom and differences in algal biomass regulation on two different substrata: fine-grained sediments and stones. The algal spring bloom was initiated when irradiance at the sediment surface exceeded 7 mol photons m^-2 d^-1 and mean water velocity was concomitantly below the threshold for bed load transport in the stream. Large inter-annual and substratum-dependent differences in peak algal biomass were observed, thus suggesting that different parameters regulate algal biomass development on the two substrata. On fine-grained sediments algal biomass development was predominantly coupled to light availability, while on stony substrata algal composition and peak biomass might be affected by invertebrate grazing.     

Organic matter availability structures microbial biomass and activity in a Mediterranean stream

1. We compared microbial biomass (bacteria, fungi, algae) and the activity of extracellular enzymes used in the decomposition of organic matter (OM) among different benthic substrata (leaves, coarse and fine substrata) over one hydrological year in a Mediterranean stream.
2. Microbial heterotrophic biomass (bacteria plus fungi) was generally higher than autotrophic biomass (algae), except during short periods of high light availability in the spring and winter. During these periods, sources of OM shifted towards autochthonous sources derived mainly from algae, which was demonstrated by high algal biomass and peptidase activity in benthic communities.
3. Heterotrophic activity peaked in the autumn. Bacterial and fungal biomass increased with the decomposition of cellulose and hemicellulose compounds from leaf material. Later, lignin decomposition was stimulated in fine (sand, gravel) and coarse (rocks, boulders and cobbles) substrata by the accumulation of fine detritus.
4. The Mediterranean summer drought provoked an earlier leaf fall. The resumption of the water flow caused the weathering of riparian soils and subsequently a large increase in dissolved organic carbon and nitrate, which led to growth of bacteria and fungi.     

Bacteria biofilm encourages algal immigration onto substrata in lotic systems

I tested the effect of the density of attached bacteria on the amount of algal immigration in the early development of a periphyton community in an artificial stream by manipulating the density of the attached bacteria. Three densities were prepared by regulation of the incubation time. A suspension of algae was added to the stream, and the degree of algal attachment to substrata was compared among the treatments. Algal immigration was proportional to the density of attached bacteria on all substrata (glass, PVC, and slate), although density differed among substrata. Analysis of covariance (dependent variable, amount of attached algae; covariate, bacterial density) showed significant relationship between amounts of attached algae and bacterial densities, but did not show significant differences in the slopes and adjusted means among substrata. When acrylic beads were added with the suspension of attached algae, significant linear correlation was obtained between the amount of attached algae and the amount of acrylic beads on the substrata. Algal immigration was due to non-selective adsorption by attached bacterial biofilms on substrata, although the extent of bacterial colonization and biofilm formation may be affected by the substrata and other environmental factors (e.g., current conditions and water temperature).     

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.     

Contribution of biofilm‐dwelling consumers to pelagic–benthic coupling in a large river

1. Over the course of this 17‐month study, we assessed the potential loss of plankton (bacteria, algae, heterotrophic flagellates) to consumers (ciliates and rotifers) within mature biofilms established on natural substrata exposed to the main current of the River Rhine (Germany). Once a month, in flow cells in a bypass system to the River Rhine, we measured the clearance rates of the biofilm‐associated consumers on the different groups within the natural plankton. 2. Ciliates were the most dominant consumers, among which planktivorous groups, particularly peritrichs and (in spring and summer) heterotrichs dominated. Consumer biomass varied with season, with the highest density occurring directly after the appearance of the phytoplankton spring peak. 3. Clearance rates on plankton ranged from 96 to 565 L m^?2 d^?1 for bacteria and 66–749 L m^?2 d^?1 for algae, with a preference for algae in summer and for bacteria in winter. This pattern coincided with seasonal changes in the structures of the grazer communities. The consumers (both ciliates and rotifers with total standing stocks ranging between 19 and 572 mg C m^?2) imported a substantial amount of organic matter (between 15 and 137 mg C m^?2 d^?1) into the biofilm. 4. These results highlight the potential importance of consumers in the biofilm as a trophic link between the plankton and the benthos, a function that has hitherto mostly been attributed to filter‐feeding bivalves. In contrast to bivalves, the biofilm‐dwelling consumers show a more dynamic response towards the plankton density and composition. Such dynamic components need to be considered when estimating total plankton consumption by the benthos.     

Differences in plating efficiency of bacteria from river epilithon sampled from upper and lower surfaces of artificial substrata

The bacterial generic composition of biofilms was investigated by determining colony-forming units on 1/2 PYG (peptone, yeast extract, and glucose medium) agar plates. The biofilms developed on the upper and lower surfaces of artificial substrata submerged in river water at Unazawa in the Tamagawa River in Tokyo, Japan. Greater variation in the number and generic composition in platable colonies was obtained on the upper surface of an artificial substratum than on the lower surface. This variation corresponded to the variation in algal biomass. Accompanying the growth of the algae, specific bacteria commensal with the algae proliferated on the upper surface. The biofilm exfoliated when the surface density of chlorophyll was 11.2 g·cm^–2. When the biofilm exfoliated, the bacterial community accompanying the algae also exfoliated, and the bacterial composition altered. The bacterial composition after exfoliation resembled that observed on the lower surface. The generic composition on the upper surface soon after it was initially submerged resembled that observed in the river water. The bacterial generic composition on the lower surface of the artificial substrata did not change greatly throughout the investigation because of the constant environmental conditions. The difference between the bacterial generic composition on the upper and lower surfaces was due to the fact that bacteria commensal with algae proliferated on the upper surface but not on the lower.     

The effect of fetch on periphyton spatial variation

The marked spatial variation in periphyton could reflect differences in exposure, grazing or substratum. To determine if any of these factors was significant, I studied the relationship between the degree of exposure to waves (measured as fetch), grazing intensity (measured as invertebrate biomass) and spatial variation in periphyton biomass in sites of similar substratum along an island in the central mesotrophic basin of Lake Memphremagog (Québec). In spring, there was a positive relationship between fetch and periphyton biomass both on stones and on artificial substrata. This effect was especially strong for diatoms, for algae between 100–1000 µm³, for planktonic forms and for filamentous and long-stalked species. In spring, water renewal encouraged growth of these forms, but the effect disappeared in summer, coincident with silica depletion. Grazers, which were not important in the spatial variation observed in spring, probably contributed to the sudden decline of periphyton in the exposed sites in summer. Observations in nearby lakes indicate that these patterns may be general.Contribution of the Lake Memphremagog Project, Limnology Research Centre.     

Nutritional value of detritus and algae in blenny territories on the Great Barrier Reef

The salariin blenny, Salarias patzneri, primarily ingests detrital aggregates and inorganic sediment, with small quantities of filamentous algae. Samples of particulate matter <125 μm and filamentous algae were collected from the territories of S. patzneri and the nutritional value of these resources assessed using organic content and protein/energy ratios calculated from protein, carbohydrate and lipid content. Samples were collected throughout the day, during both summer and winter, to enable temporal comparisons of nutritional value. The particulates <125 μm, which were predominantly amorphic detritus, accounted for 41±2% and 44±3% of the organic matter in the epilithic algal matrix (EAM) during the summer and winter, respectively, whilst filamentous algae accounted for 37±3% and 41±3% of the organic matter in the summer and winter, respectively. Carbohydrate and lipid concentration in the algal samples was significantly greater than in particulate matter <125 μm, although no significant difference was detected in protein levels of algal samples and particulates <125 μm. Consequently, the protein/energy ratio of particulates <125 μm in the summer (11±1 mg kJ⁻¹) and winter (10±1 mg kJ⁻¹) was similar to that of filamentous algae in the summer (9±1 mg kJ⁻¹) and winter (8±1 mg kJ⁻¹). These results suggest detrital aggregates are at least comparable to filamentous algae as a nutritional resource. The large contribution of high quality detritus to organic matter in the epilithic algal matrix collected from S. patzneri territories throughout the day and between seasons provides strong evidence that detritus is a valuable component of small territorial fish diets and is an integral part of coral reef food webs.     

Effect of primary producers on the heterotrophic metabolism of a stream biofilm

1. ,The influence of benthic algae on heterotrophic metabolism in a forested Mediterranean stream was investigated. Bacterial density and ectoenzymatic activities, as well as algal biomass (chlorophyll- a ) and metabolism (the rate of ¹⁴C incorporation), were measured during colonization over 60 days of artificial substrata (clay tiles) under light and dark conditions.
2. ,Chlorophyll- a and the rate of ¹⁴C incorporation were significantly higher in light-grown than in dark-grown biofilms. Bacterial density and ectoenzymatic activity (especially β-glucosidase) were also significantly higher in light-grown biofilms.
3. ,Regressions of chlorophyll- a and ¹⁴C incorporation values on the ectoenzymatic activities were significant. The slopes of regression lines obtained for dark-grown biofilms were significantly higher than those obtained for light -grown biofilms.
4. ,The differences in the slope (of the regression lines) between dark and light-grown biofilms suggest that the response of the heterotrophs is faster in biofilms with low algal biomass accrual and slows down when algal biomass is increased.
5. ,It is concluded that algal accumulation in the epilithic biofilm influences the use of organic matter by the heterotrophic community by increasing the amount of organic substrate available for bacteria.     

Successional dynamics of the phytoplankton in the lower part of the river Ebro

The temporal and spatial dynamics of phytoplankton have beenstudied in four sites located along the last 60 km of the riverEbro, over a period of 1 year. Diatoms and green algae werethe most abundant groups; blue-green algae were frequent onlyin autumn. Asterionella formosa dominated the winter phytoplanktonassemblages. In autumn, spring and early summer centric diatomswere dominant: Aulacoseira granulata (Ehr.) Simonsen in autunm;Cyclotella sp. p1., Skeletonema potamos (Weber) Hasle and StephanodisciLssp. p1. in spring. A great abundance of green algae was observedduring the summer, mainly in the lower sites. In the sites closerto the mouth, the spring maximum of centric diatoms extendedto the summer. Mainly in the downstream sites, a remarkablegrowth of Acrinocyclus normanii f. subsalsa (Juhl.-Daunf.) Hustedtand Stephanodiscus hantzschii f tenuis (Hust.) Hak. & Stoerm.was added to green algae in the late summer. As has been investigatedthrough a principal component analysis, the phytoplankton temporalsuccession and longitudinal differences between the sites maybe affected by the variations in flow and the increase of waterconductivity downstream; both factors seem to act together.The river is rather homogeneous with respect to the phytoplanktonassemblages during the winter and spring months, and from latespring to the following autumn, differences greatly increaseboth in time and downstream.     

Effects of intensity and seasonal timing of disturbances on a rocky intertidal benthic community on the southern coast of Korea

The effects of intensity and timing of disturbances on recovery of marine benthic organisms were investigated on a rocky intertidal shore in Gwangyang Bay, Korea. We hypothesized that the recovery pattern of the benthic community structure would be affected by disturbance intensity and season. Twenty-eight permanent plots were set up, with disturbance intensity (cleared plots and sterile plots) and seasonal disturbance (fall 1999 and spring 2000) incorporated into the experimental design. To monitor natural seasonal variation in benthic community abundances, we established seven permanent unmanipulated plots. Turf-forming algae were observed in the unmanipulated plots throughout the experimental period, whereas green algae and invertebrate presence varied with season. In the disturbance-intensity experiment, turf-forming and green algae were dominant in cleared plots. The highest coverage of sessile organisms was observed in sterile plots, which exhibited the highest species richness because of their relatively low macroalgal coverage. Seasonal effects of disturbance were an important factor in the recovery pattern of benthic organisms under high disturbance intensity. Coverage of green algae was higher in sterile spring plots than in sterile fall plots; this result was attributed to low spatial competition, as the disturbances occurred just before green algal blooms. On the other hand, the abundances of barnacles and bivalves were highest on sterile fall plots, as these organisms were suppressed by green algal blooms in other periods. These results indicate that the effects of disturbance intensity on benthic community recovery patterns can be influenced by season of disturbance.     

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.     

Direct and Indirect Effects of Solar Ultraviolet Radiation on Attached Bacteria and Algae in Lotic Systems

I examined the effects of solar ultraviolet radiation (UVR) on attached bacteria and algal densities in lotic systems in outdoor artificial stream apparatus. Flumes were covered with four types of film for UVR screening treatments, and attached bacterial cell densities and their temporal variations were compared between conditions excluding and including solar UVR. Attached bacterial cell densities were depressed by solar UVR, and both accrual rate and saturated density were significantly lower in the +UVR (full solar radiation) condition than in −UVR and dark conditions. Solar UVR also indirectly affected the rate of algal accrual. Microscopic direct observations of attached bacterial cell density and algae on substrata showed that solar UVR depressed the accrual of attached bacteria and consequently the frequency of sites with high bacterial cell density that can trap suspended algae in the water. The final amount of algal accrual in the +UVR condition was one-fourth of that in the −UVR condition. Therefore, the effects of solar UVR may be more serious in systems where periphyton are frequently removed by floods.     

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.     

Patterns of Biofilm Succession on a Sheltered Rocky Shore in Hong Kong

Successional patterns are dependent on the nature of the substratum, water flow, concentrations of organics as well as the availability of bacteria, algal spores and invertebrate larvae in the coastal environment. Bacteria play an especially important role in biofilm formation as they are generally the earliest colonizers. In the present study, both winter and summer biofilm succession patterns were examined on glass coverslips inverted on experimental racks attached at two tidal levels on a sheltered shore in Hong Kong. In the succession, bacteria were followed by diatoms and cyanobacteria. Encrusting algae appeared in the late stages of the experiment (day 80 in summer and day 60 in winter). Colonization by bacteria was much slower in summer and their density remained low throughout the experimental period. The first appearance of diatoms and cyanobacteria, however, was more rapid in the summer. Bacteria and diatoms on the low-shore surfaces also had a faster succession rate than on the high-shore surfaces, suggesting that desiccation/aerial temperature are the causal factors for such differences.     

Benthic algal response to N and P enrichment along a pH gradient

Nutrient enrichment and its effect on benthic algal growth, community composition, and average cell size was assessed across two sites of differing pH within a single habitat. Nutrients were added using in situ substrata, which released either N, P, or no additional nutrients (controls) at each site for 21 days. Upon collection, chlorophyll and biovolume standing stocks of the attached algal microflora were measured. Chlorophyll concentration was different among all treatments, accumulating greatest on P, followed by N, and the least on C substrata (P < 0.001) and was highest at site-2 (P < 0.001), while total algal biovolume was highest on P compared to both N and C substrata (P < 0.05) and did not vary between sites. Increased growth on P substrata was due to the enhanced biovolume of filamentous green algae, although the affected taxa varied between sites. Biovolume to cell density ratios (as a measure of average cell size) were highest on P substrata over both N-enriched and control substrata (P < 0.05) and this pattern was similar between sites. Progression towards a community composed of larger cells following P enrichment observed along this pH gradient, seems to be related to the dominance of larger celled filamentous green algae. Thus, nutrients exhibited greater control on benthic algal growth than did changes in hydrogen ion concentration.Contribution number 581, Great Lakes Environmental Research LaboratoryContribution number 581, Great Lakes Environmental Research Laboratory     

Patterns in the epilithic community of a lake littoral

1. The algae and sedentary macroinvertebrates on the upper surface of stones from the littoral of Crosemere were investigated over 13 months. This lake is one of a series of eutrophic meres of glacial origin in the English Midlands. 2. Stones were taken from shallow and deep areas in the littoral and from areas shaded by bankside trees and those away from trees. This gave four habitat types: open/shallow; open/deep; shade/shallow; shade/deep. Epilithic algae and sedentary macroinvertebrates from the upper surfaces of the stones were quantified monthly for each habitat type. 3. Chlorophyll a and ash-free dry mass both showed a strong seasonal pattern common to all habitat types, with a spring peak declining to a summer minimum, followed by a small autumn recovery, and a winter minimum. 4. In terms of percentage cover, Cladophora glomerata showed a markedly different pattern. There were strong differences between habitat types, with shaded stones from the shallows, in particular, having very sparse Cladophora cover. In the open, Cladophora cover was high in summer and low at other times. 5. The invertebrate community was dominated by retreat-dwelling larvae of the psychomyiid caddis, Tinodes waeneri, and four species of chironomids with tube-building larvae, Cricotopus sylvestris, Microtendipes pedellus, Glyptotendipes pallens and Endochironomus albipennis. For Tinodes, Cricotopus and Microtendipes, peaks of density occurred chiefly beneath trees in spring and summer. 6. The seasonal pattern of algal abundance showed little relationship with that of invertebrate biomass. The ratio of chlorophyll a to ash-free dry mass also declined in summer, despite the higher invertebrate biomass. This indicated that grazing was not the dominant factor diminishing algal abundance seasonally. It seems likely that algae were limited chiefly by physical factors, such as light and temperature, and by nutrients, particularly nitrates, which decline in summer in the epilimnion of the lake. 7. Grazing may have contributed to spatial patchiness of algae in summer, however, particularly that of Cladophora. The scarcity of Cladophora on shallow, shaded stones coincided with a high abundance of Tinodes on these stones in early summer. Riparian trees could thus have affected epilithic algae, not only by shading but also indirectly through the supply of grazers.     

Patterns of biofilm succession on a sheltered rocky shore in Hong Kong

Successional patterns are dependent on the nature of the substratum, water flow, concentrations of organics as well as the availability of bacteria, algal spores and invertebrate larvae in the coastal environment. Bacteria play an especially important role in biofilm formation as they are generally the earliest colonizers. In the present study, both winter and summer biofilm succession patterns were examined on glass coverslips inverted on experimental racks attached at two tidal levels on a sheltered shore in Hong Kong. In the succession, bacteria were followed by diatoms and cyanobacteria. Encrusting algae appeared in the late stages of the experiment (day 80 in summer and day 60 in winter). Colonization by bacteria was much slower in summer and their density remained low throughout the experimental period. The first appearance of diatoms and cyanobacteria, however, was more rapid in the summer. Bacteria and diatoms on the low-shore surfaces also had a faster succession rate than on the high-shore surfaces, suggesting that desiccation/aerial temperature are the causal factors for such differences.