A multi-lake comparison of epilithic diatom communities on natural and artificial substrates

In a multi-season, multi-lake study of epilithic diatom communities, glass slide artificial substrate samplers provided poor representation of communities on natural substrates. Percent similarities between the two communities averaged only 37 over the course of the study. Overall, natural substrates exhibited greater species richness than artificial samplers, particularly during the summer, although this difference was slight. Patterns of selection and inhibition by artificial substrates of individual diatom species varied by both season and lake. Members of the genus Cymbella, particularly C. microcephala, appeared to be the most consistently inhibited, while Achnanthes minutissima was often selected for. In spite of the great differences between the two substrate types, replicability of artificial substrates was very high, and could prove to be a more important quality in monitoring applications than accurate representation of the natural community.     

Seasonal changes in the epiphyte community of natural and artificial macrophytes in Lake Memphremagog (Que. & Vt.)

Seasonal changes in the epiphyte biomass, measured both as chlorophyll a and as cell volume, and species composition were compared on Potamogeton richardsonii and on a similar plastic plant grown together in a shallow bay of Lake Memphremagog (46°06'N, 72°16W). Both substrates exhibited two periods of high biomass during the June to September growing season; one in June, when the community was dominated by loosely attached species with a strong planktonic component (up to 37%), and one in September, when the epiphytes were characterized by species tightly attached to the leaves. Although this seasonal trend was similar, the loosely and tightly attached communities were best developed on the natural and artificial plants, respectively. The diversity of the epiphytes was significantly higher on the natural than on the artificial leaves from July on. Both the diversity differences and differences in community structure appear to be the result of the summer accumulation of CaCO₃ observed only on the upper leaf surfaces of the natural plants. Consequently, P. richardsonii appears to affect epiphyte development largely by its precipitation of CaCO₃, with no evidence for either direct inhibition or stimulation of the epiphytes by the natural plants. The reduced epiphyte biomass on growing tips was no different from that on artificial plants of the same age and exposure and is attributable to an insufficient time for colonization rather than to inhibition by the macrophyte.Contribution No. 8, Lake Memphremagog Project, Limnology Research Group     

The potential of fish production based on periphyton

Periphyton is composed of attached plant andanimal organisms embedded in amucopolysaccharide matrix. This reviewsummarizes research on periphyton-based fishproduction and on periphyton productivity andingestion by fish, and explores the potentialof developing periphyton-based aquaculture.Important systems with periphyton arebrush-parks in lagoon areas and freshwaterponds with maximum extrapolated fish productionof 8 t ha^–1 y^–1 and 7 t ha^–1y^–1, respectively. Experiments with avariety of substrates and fish species havebeen done, sometimes with supplemental feeding.In most experiments, fish production wasgreater with additional substrates compared tocontrols without substrates. Colonization ofsubstrates starts with the deposition oforganic substances and attraction of bacteria,followed by algae and invertebrates. Afterinitial colonization, biomass density increasesto a maximum when competition for light andnutrients prevents a further increase. Often,more than 50% of the periphyton ash-free drymatter is of non-algal origin. Highest biomass(dm) in natural systems ranges from 0 to 700g m^–2 and in aquaculture experiments wasaround 100 g m^–2. Highest productivity wasfound on bamboo in brush-parks (7.9 gC m^–2 d^–1) and on coral reefs (3 gC m^–2 d^–1). Inorganic and organicnutrients stimulate periphyton production.Grazing is the main factor determiningperiphyton density, while substrate type alsoaffects productivity and biomass. Better growthwas observed on natural (tree branches andbamboo) than on artifical materials (plasticand PVC). Many herbivorous and omnivorous fishcan utilize periphyton. Estimates of periphytoningestion by fish range from 0.24 to 112 mg dm(g fish)^–1 d^–1. Ingestion rates areinfluenced by temperature, fish size, fishspecies and the nutritional quality of theperiphyton. Periphyton composition is generallysimilar to that of natural feeds in fishponds,with a higher ash content due to the entrapmentof sand particles and formation of carbonates.Protein/Metabolizable Energy (P/ME) ratios ofperiphyton vary from 10 to 40 kJ g^–1.Overall assimilation efficiency of fish growingon periphyton was 20–50%. The limited work onfeed conversion ratios resulted in valuesbetween 2 and 3. A simple simulation model ofperiphyton-based fish production estimates fishproduction at approximately 2.8 t ha^–1y^–1. Together with other food resources infishponds, total fish production with thecurrent technology level is estimated at about5 t ha^–1 y^–1. Because grazingpressure is determined by fish stocking rates,productivity of periphyton is currently themain factor limiting fish production. Weconclude that periphyton can increase theproductivity and efficiency of aquaculturesystems, but more research is needed foroptimization. Areas for attention include theimplementation and control of periphytonproduction (nutrient levels, substate types andconformations), the ratio of fish to periphytonbiomass, options for utilizing periphyton inintensive aquaculture systems and with marinefish, and possibilities for periphyton-basedshrimp culture.     

A comparison of diatom community structure on natural and artificial substrata

Artificial substrata have been used in diatom studies for almost 100 years. However, concern still exists over whether diatom communities developing on artificial substrata accurately represent communities developing on natural substrata. This study compares the diatom communities colonising glass slides and clay tiles in two coastal dune lakes, and compares these communities to the naturally occurring communities in the epipelon, epilithon, and epiphyton. Both glass microslides and clay tiles, incubated for three separate periods ranging from 29 to 68 days, resulted in replicate substratum samples supporting similar diatom community compositions at each site. The degree of variation between artificial substrata communities at different sites, and between the two artificial substrata types, was generally no more than the degree of variation between communities on different types of natural substrata. Additionally, the composition of the diatom communities on the artificial substrata was representative of the community composition on the natural substrata. The effects of incubation period and siting are discussed.     

Effects of cadmium stress on periphytic diatom communities in indoor artificial streams

SUMMARY 1. Two experiments were performed with periphytic diatoms originating from the River Lot (France) and allowed to grow on clean substrata within indoor artificial streams. Three cadmium (Cd) levels (control, low Cd=10 μg L⁻¹ and high Cd=100 μg L⁻¹) were used to test the effects of Cd on (i) the settlement and development of diatom communities (Experiment 1, 4 weeks Cd exposure) and (ii) predeveloped communities grown over a 2-week period without contamination (Experiment 2, 2 weeks Cd exposure).
2. Experiment 1 revealed that growth and taxonomic composition of diatom communities clearly differed with Cd exposure. Biofilms were more adpressed to substrata under low Cd concentration, and were thinner and patchy under high Cd concentration.
3. Exposure of developed diatom communities (Experiment 2) to Cd revealed only minor variations in taxonomic composition, possibly linked to the protective role of the developed organic matrix against metal stress.
4. These results support the validity of periphytic diatom communities as indicators of metal pollution, although significant structural changes would take longer within developed communities.     

白洋淀附着藻类的初级生产力及其与水质的关系

附着藻类是湖泊中主要的生产者,尤其是草型湖泊。但与浮游藻类相比,针对附着藻类初级生产的研究还相对较少。采用原位调查与实验模拟相结合的方法测定2014—2015年间白洋淀附植藻类和附泥藻类的现存量和初级生产力,并对附着藻类初级生产与白洋淀水体理化参数的关系进行分析。结果表明,不同采样季节的附植藻类和附泥藻类的叶绿素a分别为34.83—245.22μg/cm~2和26.08—297.40μg/cm~2,无灰干重分别为0.46—5.21g/m~2和0.61—5.81g/m~2。两种附着藻类的生物量都在8月最高,4月和11月最低。空间分布上,南刘庄、府河入口的附着藻类生物量显著高于采蒲台和枣林庄。白洋淀附植藻类和附泥藻类的年均总初级生产分别为494.20mg C m~(-2)d~(-1)和474.45mg C m~(-2)d~(-1),呼吸速率为522.63mg C m~(-2)d~(-1)和508.98mg C m~(-2)d~(-1),净初级生产为-28.44mg C m~(-2)d~(-1)和-34.52mg C m~(-2)d~(-1)。白洋淀附着藻类初级生产力具有明显的时空分布规律,8月最高,6月次之,4月和11月最低,空间分布呈自西向东递减的趋势,在府河入淀口和南刘庄处最高,枣林庄和采蒲台最低。水质较好的区域的净初级生产力为正值,表明这些区域附着藻类以自养型群落为主,水质较差区域的净初级生产力为负值,则该区域以异养型群落为主。运用冗余分析法(RDA)探讨附着藻类与水质因子之间的关系,并采用向前引入法对水质因子进行逐步筛选,Monte Carlo置换检验结果显示,总磷、浮游植物叶绿素a、高锰酸盐指数、氨氮、水温、透明度、溶解氧和氮磷比是影响附着藻类生物量和初级生产的关键水质因子。附着藻类的总初级生产与水体富营养化程度呈正相关关系。     

Effects of nutrients and light on periphyton biomass and nitrogen uptake in Mediterranean streams with contrasting land uses

1. Nutrient diffusing substrata (NDS) were used to determine the relative importance of nutrients and light as potential limiting factors of periphyton biomass and nitrogen (N) uptake in Mediterranean streams subjected to different human impacts. The nutrients examined were phosphorus (P) and N, and we also further differentiated between the response of periphyton communities to N species (i.e. NO₃‐N and NH₄‐N). To examine the effect of light and nutrients on periphyton biomass, chlorophyll a accrual rates on NDS located at open and closed canopy sites were compared. The effect of nutrient availability on periphyton uptake was measured by ¹⁵N changes on the NDS after NO₃‐¹⁵N short‐term nutrient additions. 2. Results show that light was the main factor affecting algal biomass in the study streams. Algal biomass was in general higher at open than at closed canopy sites. Nutrient availability, as simulated with the NDS experiments, did not enhance algal biomass accrual in either of the 2 light conditions. 3. In the control treatments (i.e. ambient concentrations), periphyton NO₃‐N uptake rates increased and C : N molar ratios decreased consistently with increases in N availability across streams. NO₃‐N uptake rates were altered when ambient N concentrations were increased artificially in the N amended NDS. Periphyton assemblages growing on N enriched substrata seemed to preferentially take up N diffusing from the substratum rather than N from the water column. This response differed among streams, and depended on ambient N availability. 4. Periphyton biomass was not significantly different between substrata exposed to the two forms of available N sources. Nonetheless, we found differences in the effects of both N sources on the uptake of N from the water column. NH₄‐N seemed to be the preferred source of N for periphyton growing on NDS. 5. Results suggest that the effect of riparian zones on light availability, although seldom considered by water managers, may be more important than nutrients in controlling eutrophication effects derived from human activities. Finally, our results confirm that not only increases in concentration, but also stoichiometric imbalances should be considered when examining N retention in human altered streams.     

Effects of copper on periphyton communities assessed in situ using chemical-diffusing substrates

Chemical-diffusing substrates were designed to allow delivery of toxicants to mature periphyton communities under natural conditions without contaminating the surrounding environment. Artificial stream validation studies were conducted in which the effects of substrate-released copper (Cu) on periphyton communities were compared to those generated in a more conventional manner (via water column additions). Effects of copper on the following community parameters were assessed: total community biomass (measured as ash-free dry mass), relative chlorophyll a (chl a and adenosine triphosphate contents, and relative biomass of heterotrophic bacteria. Exposure of more laboratory periphyton communities to substrate-released Cu generated dose-response relationships and recovery models that were indistinguishable from those generated by the conventional route of exposure. The results of this study demonstrate the utility of chemical-diffusing substrates in field validations of laboratory toxicity tests and in investigations of the effects of stress history on periphyton tolerance to toxicants. This revised version was published online in September 2006 with corrections to the Cover Date.     

The independent and interactive effects of snail grazing and nutrient enrichment on structuring periphyton communities

We investigated the independent and interactive effects of nutrient enrichment and snail grazing on structuring periphyton communities in a northern temperate lake. Nutrient releasing substrates and grazer enclosures were used to simultaneously manipulate nutrient availability and herbivory. Periphyton was allowed 18 days to accrue before grazers (Elimia livescens = Goniobasis livescens) were introduced.Addition of nitrogen and phosphorus caused a significant increase in biovolume (p < 0.001), whereas grazing had no significant effect on biovolume but resulted in a shift in species composition. Four taxa were largely responsible for the increase in biovolume on the nutrient enriched substrates: Oedogonium sp, Stigeoclonium tenue, Navicula radiosa var. radiosa and Navicula radiosa var. tenella. By the 28th day, nutrient enrichment caused a shift from a community dominated by diatoms (Bacillariophyceae) to a community dominated by green algae (Chlorophyceae). Blue green algae (Myxophyceae) maintained an equal proportion in high and low-nutrient regimes.Grazing had a more pronounced effect on altering community composition on the nutrient enriched substrates than on the unenriched substrates. Grazing caused a decrease in diversity and an increase in dominance by green algae on the nutrient enriched substrates. The relative biovolume of green algae increased from 64% to 93% on grazed substrates, due to the significant increase in relative abundance of Stigeoclonium tenue. This taxon has both prostate basal cells and erect filamentous cells. The ratio of basal: filamentous cells increased from 4.7 to 5.2 with grazing, suggesting that the heretotrichous growth form of Stigeoclonium tenue is adapted to grazing by virtue of the basal cells which are able to adhere to the substratum and resist being grazed.     

Fractal geometry and root system structures of heterogeneous plant communities

Above-ground plant growth is widely known in terms of structural diversity. Likewise, the below-ground growth presents a mosaic of heterogeneous structures of differing complexity. In this study, root system structures of heterogeneous plant communities were recorded as integral systems by using the trench profile method. Fractal dimensions of the root images were calculated from image files by the box-counting method. This method allows the structural complexity of such associations to be compared between plant communities, with regard to their potentials for soil resource acquisition and utilization. Distinct and partly significant differences are found (fractal dimension between 1.46±0.09 and 1.71±0.05) in the below-ground structural complexity of plant communities, belonging to different biotope types. The size of the heterogeneous plant community to be examined has an crucial influence on the fractal dimension of the root system structures. The structural heterogeneity becomes particularly evident (fractal dimensions between 1.32 and 1.77) when analysing many small units of a complex root system association. In larger plant communities, a broad variety of below-ground structures is recorded in its entirety, integrating the specific features of single sub-structures. In that way, extreme fractal dimensions are lost and the diversity decreases. Therefore, the analysis of larger units of root system associations provides a general knowledge of the complexity of root system structures for heterogeneous plant communities.     

Plant community dynamics in a chain of lakes: principal factors in the decline of rooted macrophytes with eutrophication

Shoe Lake and East Graham Lake, part of a small chain of lakes in southeastern Michigan, USA, differ in nutrient loading and in the structure and productivity of their aquatic plant communities. A comparative study of species frequency and biomass distributions, nutrient contents, and responses to experimental nutrient enrichment and shading, was conducted to determine the principal factors controlling the macrophyte dynamics. A central objective was to address the question of why rooted macrophyte growth declines with eutrophication, and to test existing models designed to explain this phenomenon. In the more eutrophic Shoe Lake, diversity and productivity of rooted macrophytes were relatively low, restricted primarily by combined shading of phytoplankton, periphyton, and non-rooted macrophytes (principally Ceratophyllum demersum, along with Utricularia vulgaris and Cladophora fracta). In the less eutrophic East Graham Lake, lower nitrogen availability restricted the growth of all of these shading components, resulting in clearer water and higher productivity and diversity of rooted macrophytes. The macrophytes did not allelopathically suppress the phytoplankton in East Graham Lake. The results supported a direct relationship between nutrient loading, increasing growth of phytoplankton, periphyton and non-rooted macrophytes, and decline of rooted macrophytes.     

Structural characteristics of algal communities in thermally altered artificial streams

Algal community structure on natural substrates of thermally altered artificial streams was studied for one year. The streams were fed by a natural blackwater stream near Aiken, South Carolina. Temperature-related shifts in the abundance of major species were demonstrated. Red algae were absent from a stream heated 12·5°C above ambient, but remained abundant in streams heated 7·5°C or less. Water temperatures above 30°C produced blue-green algal dominance and eliminated several indigenous species. Substrate specificity was exhibited by all major taxa. Several taxa were abundant only on the bottom sediments and some showed a distinct affinity for either sand or silt, the principal bottom types.     

Impact of nutrient enrichment and alkalinization on periphyton communities in the New Jersey Pine Barrens

The impact of residential and agricultural development on stream periphyton communities in the New Jersey Pine Barrens was examined by comparison with communities in undeveloped areas. Watershed disturbance resulted in stream water primarily characterized by greatly elevated pH levels and nitrate concentrations. A total of 53 periphyton species were encountered in bimonthly samples over a one year period in the three disturbed and three undisturbed study streams. Species richness was significantly greater in the disturbed streams based on three criteria: the average number of species per stream on each sampling occasion (disturbed = 6.3; undisturbed = 4.9), the average number of species per stream for the entire year (disturbed = 19.3; undisturbed = 16.0), and the total number of species found in streams within a type (disturbed = 40; undisturbed = 31). Species composition also changed significantly as the result of disturbance. There appeared to be replacement of species characteristic of undisturbed Pine Barrens streams with species peripheral to the region. The expected effects of both elevated pH and nitrate were consistent with these results.     

Stable nitrogen isotope ratios of macrophytes and associated periphyton along a nitrate gradient in two subtropical, spring-fed streams

1. An increase in human population and associated changes in land use have caused an increase in groundwater nitrate concentrations throughout central Florida. Within the region, this nitrate‐laden groundwater returns to the surface via numerous large springs that serve as the origin of flow for many coastal streams and rivers. These rivers can exhibit strong nitrate gradients because of the high nutrient uptake potential of the rivers. 2. We hypothesised that downstream declines in nitrate concentrations would be manifested spatially as increases in the δ¹⁵N of the residual pool of nitrate, macrophytes and periphyton as a consequence of isotopic fractionation associated with preferential use of ¹⁴NO₃⁻. This hypothesis was tested in two spring‐fed river systems, the Chassahowitzka and Homosassa rivers, along Florida's central Gulf of Mexico coast. 3. In general, δ¹⁵N values of nitrate, macrophytes and periphyton increased with decreasing fraction of nitrate remaining in each of the two study systems. The fractionation associated with nitrate uptake by macrophytes and associated periphyton was determined from the relationship between δ¹⁵N of both constituents of the macrophyte community and the fraction of nitrate removed from the system. Values for fractionation by macrophytes and periphyton ranged from 1.9‰ to 3.6‰ and from 0.7‰ to 2.5‰, respectively.     

Impact of the Tihange nuclear power plant on the periphyton and the phytoplankton of the Meuse River (Belgium)

A survey was made from 1976 to 1981 of the possible effects of Tihange nuclear power plant on the Meuse river. Phytoplankton and periphyton were affected only during low water flow. Both decreases and increases of biomass were recorded for phytoplankton but, for periphyton, only increases. The structure of diatom communities is affected downstream of the plant during low water flow, because sensitive species are replaced by more tolerant ones. In the reach downstream of the plant, the maximum observed temperature increase is 4.2 °C and the maximum decrease in dissolved oxygen is 15%. Except for these two parameters immediately below the plant, any changes observed cannot be attributed solely to the nuclear power plant, but rather to the combination of thermal effects with effluents from other industries which together contribute to the more significant changes observed further downstream.     

Comparisons among colonization of artificial substratum types and natural substratum by benthic macroinvertebrates

A field study was designed to concurrently evaluate differences in colonization by benthic macroinvertebrates on a range of artificial substratum types (single particles of natural rock or clay brick and baskets of natural substratum) after three colonization periods (1, 8 and 29 days). Fauna on the artificial substrata were compared to natural substratum and the effect of natural epilithic cover on colonization by zoobenthos was determined. Densities of total number of organisms and the seven most abundant taxa, total number of taxa and quantity of organic material were greater on the natural substratum than on the artificial substratum types. Relative abundances of taxa on pairs of the artificial substratum types, unlike pairs of each artificial substratum type and the natural substratum, were statistically correlated. Among the artificial substratum types densities of total number of organisms and about one-half of the most abundant taxa, total number of taxa and quantity of organic material were greatest in the substratum baskets. Natural epilithic cover on the single rock particles and substratum baskets affected the densities of total number of organisms and two of the seven most abundant taxa. These taxonomic groups were at approximately two to six-fold greater densities on the substrata with fine sediment. consistent patterns in densities of the zoobenthos on the substrata were found after each colonization period. In our study all measures of the macroinvertebrate assemblages (densities of each taxon, total number of organisms, total number of taxa and relative abundances of taxa), with few exceptions, were different between each artificial substratum type and natural substratum. This result showed the abundance and composition of the macroinvertebrate fauna on artificial substratum types were different from the natural substratum. Therefore, the choice of using artificial substrata instead of direct sampling of the natural substratum should be carefully made. Among the artificial substratum types relative abundances of taxa were similar on the single substratum particles and substratum baskets indicating single particles instead of baskets might be used to sample the zoobenthos. Investigators should consider the potential effect of the natural epilithic cover of substratum particles on colonization by zoobenthos when choosing the type of artificial substratum.     

Effects of current on the development of loosely and tightly attached layers in periphyton communities

The growth rate and peak biomass of loosely and tightly attached layers in periphyton communities were measured in slow (39 cm s^?1), moderate (67 cm s^?1) and fast (137 cms^?1) current regimens using experimental channels. The peak biomasses of the loosely and tightly attached layers were highest in the moderate and fast current regimens, respectively. The growth rate of the loosely attached layer was lower in the fast current regimen than in the other current regimens, however, there was no significant difference in the growth rates of the tightly attached layers. These results indicate that current restricts the biomass accumulation of the loosely attached layer in the slow and fast current regimens, and that biomass accumulation is highest in the moderate current regimen, while in the tightly attached layer the fast current regimen led to a maximization of the biomass.     

Effects of aquatic macrophytes on water quality and phytoplankton communities in shallow lakes

We investigated aquatic macrophytes, water quality, and phytoplankton biomass and species composition in three shallow lakes with different levels of vegetation cover and nutrient concentration in Kushiro Moor, during August 2000. Trapa japonica can live in a wide range of nutrient levels. This species forms an environment with a steeper extinction of light, higher concentrations of dissolved organic carbon (DOC), lower concentrations of dissolved oxygen (DO) near the bottom, and lower concentrations of nitrate+nitrite and soluble reactive phosphorus (SRP) than other vegetation types. The pH was much higher in a Polygonum amphibium community, and the DO near the bottom did not decrease compared to a T.japonica community in the summer. The relationship between chlorophyll a and the limiting nutrient (total phosphorus (TP) when total nitrogen (TN):TPis 10 and TN/10 when TN:TP is <10) significantly differed between lakes with and without submerged vegetation. The chlorophyll a concentrations at a given nutrient level were significantly lower in water with submerged macrophytes than in water without them. Correspondence analysis showed that the difference in phytoplankton community structure across sites was largely due to the presence or absence of submerged macrophytes, and the ordination of phytoplankton species in the lakes with submerged macrophytes is best explained by environmental gradients of TN, chlorophyll, pH and SRP.     

Effects of the water discharge on periphyton abundance and diversity in a large river (River Danube,Hungary)

This paper examines the relevance of intermediate disturbance theory in the context of in a large river the algal flora attaching to artificial substrata. It was observed that the theory is in good agreement with most of the communities analysed, however, with different frequency intervals compared to phytoplankton. Floods provided the most significant disturbances. As a rough generalization, we can say that in case of floods following each other in 8–14 days, respectively, the disturbance is of medium frequency. In case of 8 days, high frequency is more typical while in case of 14 days, low frequency is more characteristic. Owing to the fact that riverine periphytic algal organisms are attached in different ways and with differing efficiencies, the changes in the total mass occurring as a result of changes in water discharge lead to changes in diversity. The analysis of individual numbers can help in investigating the intensity of the disturbing effect on the algal communities. Negative correlation was found between water discharge and numbers of individuals, that is, the numbers of algae increase with low water discharge and decrease with high water level in the river. This second observation is valid only within a certain range of discharges (in this case up to 100 m³ s^–1), which will be specific to particular rivers and individual reaches. Below the upper limit of the range, the increased water flows carry more nutrients to the organisms but with no severe wash-off effect. Beyond the upper limit (in this case in range 100–400 m³ s^–1) the numbers of individuals is roughly in inverse proportion to the wash-off effect of the current, and the community is in equilibrium. In cases when discharge surpasses 400 m³ s^–1 decrease in individual numbers is general. We suggest that in the environments similar to the study area, the periphyton of the artificial substrata (e.g., pontoons) undergo similar processes. In respect of the periphyton formed on the surface of the riverbeds in large rivers with considerable changes in water level, a more complex system can be supposed where part of the periphyton is periodically dried and flooded alternately.     

A Comparison of Phytoplankton and Algae Growing on Plexiglas Slides in an Oligotrophic Lake

This study compares the composition and dynamics of phytoplankton communities and of periphyton communities growing on Plexiglas slides in a temperate oligotrophic lake. Ordination methods were used in the comparisons, which covered a three-month period during the summer. Plankton and periphyton communities differed considerably, and shared only six common taxa. An ephemeral Dinobryon pulse occurred simultaneously in both communities, and represented the largest source of variability in community composition. Recovery from the pulse was faster in the phytoplankton than in the periphyton. Attachment to substrata, through its effect on cell turnover times and nutrient cycling, is hypothesized to provide a temporal refuge for plankton species during periods of unfavorable environmental conditions.