Periphytic Algal Community Research Articles

Differences in seasonal dynamics, ecological driving factors and assembly mechanisms of planktonic and periphytic algae in the highly urban Fenhe River

BackgroundAlgae play important roles in urban river ecosystems and are the cornerstones of most water quality monitoring programs. Thus, a better understanding of algal community dynamics is needed to support sustainable management of water resources in urban rivers.ResultsIn this study, we quantified the seasonal variations in planktonic and periphytic algal community structure in the highly urban Fenhe River and identified environmental factors affecting algal community structure and diversity. We monitored planktonic (drifting) and periphytic (attached) algal communities in the Taiyuan section of the Fenhe River over one year. The results indicated that Cyanophyta was the dominant phylum in both communities, followed by Bacillariophyta and Chlorophyta. Significant differences were observed in the composition of the planktonic and periphytic algal communities. In particular, the periphytic algal community was more diverse than the planktonic community. Water temperature and pH were the main environmental factors affecting planktonic and periphytic algal community structure, respectively, while nutrients were the most significant factor affecting planktonic and periphytic algal diversity. Ecological modeling indicated that the variations in the algal communities of the Fenhe River are mainly driven by stochastic processes. A co-occurrence network developed for the communities displayed positive interactions between the planktonic and periphytic algae.ConclusionsThese findings deepen our understanding of the seasonal interaction between planktonic and periphytic algae and the driving factors affecting community structure in the Fenhe River. They also provide a theoretical basis for the managing and protecting water resources in urban river ecosystems.

The effect of ship-induced wave trains on periphytic algal communities in the littoral zone of a large regulated river (River Danube, Austria)

Riverine ecosystems are profoundly influenced by hydrological dynamics and natural flow regimes, which dictate the temporal variability of water levels and the amplitude of fluctuations. Human activities, particularly navigation and hydropower generation, have significantly altered these natural patterns, leading to detrimental impacts on the physical, chemical, and biological integrity of river ecosystems. The littoral zone, in particular, is highly susceptible to anthropogenic disturbances, experiencing disruptions in biological activity and biogeochemical processes. This study evaluates the effects of ship-induced wave trains on the structural and functional properties of periphytic algal communities in a regulated river environment. Using data from the Austrian Danube River, periphytic algae's immediate and long-term responses to wave events generated by different types of ships were investigated. Immediate reactions of periphytic algae to wave trains were characterized by reductions in the effective quantum yield of PS II, indicating stress-induced down-regulation of photosystem II photochemistry. Abrasion and remobilization of periphytic algae due to wave action led to increased resuspension of chlorophyll-a into the water column. Furthermore, ship-induced wave trains influenced the pigment composition of periphyton, with photoprotective mechanisms being activated in response to fluctuating light conditions. Long-term effects of wave impact on periphytic algae biomass varied depending on water depth and exposure to aerial stress. While wave action mitigated desiccation stress in shallow areas, it resulted in biomass reduction and alterations in community composition in deeper zones. Notably, the occurrence of diatoms decreased in wave-impacted areas, potentially shifting the community towards Chlorophyceae dominance. Overall, this study underscores the complexity of ship-induced wave impacts on riverine ecosystems and highlights the importance of considering both immediate and long-term responses of periphytic algal communities. Understanding these dynamics is crucial for informing sustainable management strategies aimed at mitigating the adverse effects of navigation activities on riverine biodiversity and ecosystem functioning.

Competition in the Periphytic Algal Community during the Colonization Process: Evidence from the World's Largest Water Diversion Project.

Periphytic algal colonization is common in aquatic systems, but its interspecific competition remains poorly understood. In order to fill the gap, the process of periphytic algal colonization in the Middle Route of the South to North Water Diversion Project was studied. The results showed that the process was divided into three stages: the initial colonization stage (T1, 3-6 days), community formation stage (T2, 12-18 days) and primary succession stage (T3, 24-27 days). In T1, the dominant species were Diatoma vulgaris (Bory), Navicula phyllepta (Kützing) and Fragilaria amphicephaloides (Lange-Bertalot) belonging to Heterokontophyta; these species boasted wide niche widths (NWs), low niche overlap (NO) and low ecological response rates (ERRs). In T2, the dominant species were Diatoma vulgaris, Cymbella affinis (Kützing), Navicula phyllepta, Fragilaria amphicephaloides, Gogorevia exilis (Kützing), Melosira varians (C.Agardh), Phormidium willei (N.L.Gardner) and Cladophora rivularis (Kuntze). These species displayed wider NWs, lower NO, and lower ERRs than those in T1. In T3, the dominant species were Diatoma vulgaris, Cymbella affinis, Navicula phyllepta, Fragilaria amphicephaloides, Achnanthes exigu (Grunow), etc. Among them, Heterokontophyta such as Diatoma vulgaris and Cymbella affinis had a competitive advantage based on NWs and ERRs. Cyanobacteria like Phormidium willei lost their dominant status due to the narrower NW and the increased NO. It could be concluded the interspecific competition became fiercer and shaped the colonization process; this study will be helpful in understanding the colonization of periphytic algal communities.

Longitudinal effects of land-cover transitions on the periphyton community of a tropical stream

Our study aims to investigate the longitudinal effects of two land-cover transitions on the periphytic algal community. We utilized datasets from three different studies conducted over a 5-year interval in a tropical headwater stream. The studied stream traverses two abrupt adjacent transitions from an upstream forest to a pasture and back to a downstream forest remnant. We performed a high-spatial resolution sampling and used generalized additive models (GAMs) to capture the non-linear gradient response of algal metrics to distance from land-cover transitions. Algal biomass presented a lagged response to increased light availability along the pasture section and decreased along a shorter distance in the downstream forest. Most algal metrics presented a lagged response to transitions, with chlorophyll-a taking up to 375 m to reach the maximum values inside the pasture and up to 300 m to return to reference conditions inside the downstream forest. In the downstream forested section, diatom richness and abundance were similar to the upstream forested section but did not return to reference conditions. The results were consistent across years. Our results indicate that, while riparian forest remnants can play an important role in buffering impacts related to land-cover changes in low order streams, both the magnitude and directionality of these effects might be influenced by longitudinal effects caused by the flow of water. Riparian forest remnants can have a longitudinal effect in stream conditions, influencing environmental characteristics even over non-forested reaches, to where the forest conditions can be propagated downstream by the flow of water.

Species Composition and Association of Periphyton on Ceratophyllum demersum L. and Vallisneria spiralis L. in Two Freshwater Ponds of Kavaledurga Fort, Central Western Ghats, Karnataka, India

The aim of the current research was to ascertain how the periphytic algae's species composition and diversity varied seasonally and how those changes related to the physico-chemical elements of two submerged macrophytes (<i>Ceratophyllum demersum, Vallisneria spiralis</i>) in the ponds of Kavaledurga Fort. Periphytic algal communities were highly diversified totalling 70 species, and limnological variables were measured in pond water samples over the three seasons of the study period from September 2020 to May 2021. Members of <i>Zygnematophyceae (n = </i>57) were found in water bodies with a high diversity and covering of macrophytes. <i>Ceratophyllum demersum (n =</i> 64) was an algae rich macrophyte when compared to <i>V. spiralis (n = </i>25). The main determining factors for this community variability were elevations in temperature, pH, DO, electrical conductivity and nutrients. The species richness and evenness of periphytic algae over the macrophytes were disclosed by diversity indices. From the similarity index, the results indicated a slight difference in populations of periphytic micro-algae between the studied ponds.

Spatial pattern of periphytic algae communities in major tributaries of the upper Jinsha River and the environmental drivers

Spatial pattern of periphytic algae communities in major tributaries of the upper Jinsha River and the environmental drivers

Response of periphytic algae community structure characteristics to hydrodynamic conditions in an open channel

Response of periphytic algae community structure characteristics to hydrodynamic conditions in an open channel

<em>In situ</em> effects of arsenic, aluminium and chromium stresses on algal periphyton of the river Ganga at Varanasi, India

In situ effect of metal stress on periphytic algal communities of a river was studied using chemical diffusing substrates. The metal stress caused the inhibition of periphytic biomass in a concentration-dependent manner. The study indicated differential response of various periphytic groups to different metal treatments. Diatoms exhibited tolerance against arsenic (As) and aluminium (Al) treatment but displayed sensitivity against chromium (Cr) treatment. An increased abundance of cyanobacteria was noteworthy in Cr enrichment, but Al and As were hazardous to these organisms. The relative abundance of green algae also increased in all three test metals. The metal stress lowered the species richness and diversity of periphytic algae, apparently due to the elimination of some of the sensitive species followed by an increased abundance of tolerant forms. Periphytic taxa tolerant to one metal were not necessarily tolerant to other metals or metalloids, and vice versa. The metal-induced changes in algal community composition will lead to severe ecological consequences by affecting biological diversity and in turn productivity of aquatic systems. Since algae occupy the aquatic food web base, any harmful effect on these organisms would have repercussions at higher trophic levels. Thus, it seems urgent to incorporate biomonitoring practices and chemical analysis to monitor the river Ganga's ecological health.

Dynamic Characteristics of Periphytic Algae Communities on Different Substrates and the Host Response in Subtropical-Urban-Landscape Lakes

Outbreaks of periphytic algae, including filamentous algae, have been observed after submerged macrophyte restoration and are common in early stages. Dynamic changes in the periphytic algae community on Vallisneria natans and artificial V. natans were investigated in situ, and their characteristics were compared on the two substrates. The results showed that more periphytic algae species occurred on V. natans (77 taxa) than on artificial V. natans (66 taxa) (F = 2.089, p = 0.047). The cell density and chlorophyll a (Chl. a) content of periphytic algae were 3.42–202.62-fold and 2.07–15.50-fold higher on the artificial substrate than on V. natans, respectively. Except for Lyngbya perelagans (i.e., the only common dominant periphytic algae species on the two substrates), the dominant species on V. natans were Cocconeis placentula and Ulothrix tenerrima, while those on the artificial substrate were Stigeoclonium aestrivale, Oscillatoria tenuis and Achnanthes minutissima. The cell density of periphytic algae was significantly affected by the total phosphorus (TP) and NO3−-N and electric conductivity on V. natans, and by TP and NH4+-N on artificial V. natans. The malondialdehyde content of V. natans was significantly correlated with the periphytic algae biomass. V. natans was more affected by periphytic algae during its slow-growing period, and the contribution order of stress to V. natans was diatoms > cyanobacteria > green algae. Our findings might contribute to the understanding the effect of substrate specificity on periphytic algae communities, and have important implications for the restoration of submerged plants in eutrophic lakes.

Functional Diversity and Primary Production Predict Future Patterns of Periphyton Productivity after Species Extinction

Understanding of the trait-based ecology of the periphytic algal community has increased in the last decade. However, the relationship between their functional diversity and ecosystem functions, such as primary production, has been speculated on, but yet not proven. Human impacts promote changes in biotic communities leading to a risk of extinction, with consequences for the functioning of aquatic ecosystems. In this study, we unraveled the associations between the taxonomic and functional components of periphytic algal diversity, stream eutrophication, and productivity patterns. Furthermore, we simulated future patterns of species extinction to predict how productivity may change when facing extinction. Primary production and taxonomic and functional diversity of the periphytic algal communities were estimated in five streams across a trophic gradient in the Ave River basin (northwest Portugal). Our results demonstrated that eutrophication led to a unimodal pattern of taxonomic diversity, while functional diversity tended to increase with increasing eutrophication. We found that only functional diversity had a positive association with primary production. The extinction estimations indicated that almost all species found in our study were at high extinction risk. When we spatially scaled our extinction simulations, we found poor-productive streams after the extirpation of a few species. However, at the regional scale, the ecosystem supports the extinction of at least 40% of species before turning into a poor-productive system. Intermediate levels of disturbance are probably beneficial for the diversity of periphytic algal communities, to a certain extent. Moreover, functionally diverse communities were more productive, and the alleged future extinction of species is likely to lead to poor-productive streams if regionally focused conservation initiatives are not implemented. We recommend that, using simulations of functional extinction, it is possible to infer how the loss of these microorganisms could alter ecosystem functioning, to better predict human impacts on aquatic ecosystems.

Community Composition and Assessment of the Aquatic Ecosystem of Periphytic Algae in the Yangtze River Basin

To explore the periphytic algae community structure in the Yangtze River basin, samples were collected from 130 sampling sites, including the source to the estuary along the mainstream of the Yangtze River, eight primary tributaries, and the tributary of the Three Gorges area. The periphytic algae densities of different areas in the mainstream of the Yangtze River ranked from high to low were the upstream area, source area, middle and lower area, and the Jinsha River. The high periphytic algae density in the upstream area was associated with the shift in nutrition level, and the high periphytic algae density in the source area was associated with human activity. The spatial pattern of the periphytic algae community in the whole main stream from west to east presented the alternating dominance of Bacillariophyta and Cyanophyta; the Bacillariophyta (Navicula) had a competitive advantage in the main stream, and the distribution of the periphytic algae community was driven by total nitrogen, total phosphorus, and pH. For the tributary of the Yangtze River, the periphytic algae density in the Three Gorges tributary area was far higher than those in the eight primary tributaries; the periphytic algae community was dominated by Cyanophyta (Lyngbya), which had a competitive advantage in the tributaries of the Yangtze River. The distribution of the periphytic community was driven by dissolved oxygen and pH. According to the diversity analysis and assessment, the periphytic algae community in the source area showed lower species richness and higher evenness, thus leading to a high α-diversity and good assessment result (mesosaprobic zone). The middle and lower reaches of the Yangtze River also showed the same assessment result, the mesosaprobic zone. However, the community evenness of the middle and lower reaches was significantly lower than that of the source area, thus making the middle and lower reaches of the Yangtze River have a significantly lower α-diversity than that of the source area. All areas of the Yangtze River showed good water quality assessment; however, different areas had different WQI index numbers, and the assessment results of the WQI index were inconsistent with the results of the aquatic assessment. Therefore, a comprehensive assessment of aquatic ecosystem health should use both aquatic assessments and water quality assessments.

Environmental Drivers and Aquatic Ecosystem Assessment of Periphytic Algae at Inflow Rivers in Six Lakes over the Yangtze River Basin

Periphytic algae is frequently utilized as a health indicator for ecosystems. Many research studies have been conducted in China on the periphytic algae community, but none has compared the periphytic algae community structure at inflow rivers among different lakes in the Yangtze river basin. The periphytic algae were investigated at 94 sites in inflow rivers of Dianchi Lake, Danjiangkou Reservoir, Dongtinghu Lake, Poyanghu Lake, Chaohu Lake, and Taihu Lake. Based on microscopic research, eight phyla and 126 genera of periphytic algae were found in the inflow river of six lakes, with Cyanobacteria and Bacillariophyta dominating. The CCA (Canonical Correspondence Analysis) was used to analyze the association between the periphytic algae community and environmental factors in the inflow river of six lakes, and the LefSe (Linear discriminant analysis effect size) analysis was used to find enriched species in the inflow river of six lakes. We discovered that TN (total nitrogen) and TP (total phosphorus) were the driving environment variables at the basin scale based on the combined results of the CCA and the Mantel Test. The TITAN (Threshold Indicator Taxa Analysis) analysis also revealed the indicator species and their TN and TP concentration thresholds. Finally, we assessed the ecosystem health of the inflow river at six lakes; biotic and abiotic indices yielded conflicting results, but utilizing both indices to assess ecosystem health using the Random Forest algorithm will yield objective and comprehensive results.

Structural Characteristics of Periphytic Algal Community and Its Relationship with Environmental Factors in the Taiyuan Region of the Fenhe River

In order to explore the characteristics of the periphytic algae community structure and its relationship with environmental factors in the Taiyuan region of the Fenhe River, a total of six sampling sites were investigated in July and December 2021. The effects of water quality status and environmental factors at each sampling point on the community structure of epiphytes were detected. The results showed that a total of 7 phyla and 54 genera of periphytic algae were identified in the Taiyuan region of the Fenhe River, and the species composition was mainly Bacillariophyta, Cyanophyta, and Chlorophyta. According to the analysis results of the biodiversity index, the water body of the Taiyuan region of the Fenhe River is in a state of moderate pollution. The correlation analysis between the epiphytic algae and environmental factors showed that the cell density of algae was significantly correlated with dissolved oxygen (DO), phosphate (PO43−-P), chemical oxygen demand (COD), total phosphorus (TP), and transparency (SD) in the wet season. The algal cell density in the dry season was significantly correlated with water temperature (WT), TP, PO43−-P, and COD. According to the redundancy analysis, the community distribution of the epiphytic algae in the Taiyuan region of the Fenhe River was closely related to physical and chemical factors such as COD, nitrate nitrogen (NO−3 -N), WT, dissolved organic carbon (DOC), total nitrogen (TN), and TP, and COD is the main environmental factor driving the change in the community distribution of the periphytic algae in the wet season. TN is the main control factor driving the change in the biological community distribution of periphytic algae in the dry season.

Environmental Factors Drive Periphytic Algal Community Assembly in the Largest Long-Distance Water Diversion Channel

Periphytic algae exist widely in different waters. However, little is known about periphytic algae in long-distance water diversion channels across watersheds. We investigated the periphytic algae and the environmental factors at twenty sampling sites in the middle route of the South-to-North Water Diversion Project (MRP). The dominant species were Desmodesmus intermedius (Hegewald), Calothrix thermalis (Bornet & Flahault), Calothrix parietina (Bornet & Flahault) and Leptolyngbya benthonica (Anagnostidis) (dominance > 0.02) as measured in a whole year. Habitat heterogeneity in the MRP led to lower spatial heterogeneity and higher temporal heterogeneity of the periphytic algal community. Stochastic processes are the major process in periphytic community assembly. In deterministic processes, homogeneous selection had the major role in structuring the periphytic community, whereas the role of heterogeneous selection was less significant. In stochastic processes, dispersal limitations had the major role in structuring the periphytic community, whereas the role of homogenizing dispersal and drift were less significant. The variation in total nitrogen and total phosphorus promoted more stochastic processes (−1.96 < βNTI < 1.96). The variations in water temperature and water velocity promoted more heterogeneous selection (βNTI > 1.96). In integrating all of this empirical evidence, we explore the role of environmental factors in the action of ecological processes shaping thecommunity assembly of the periphytic algal community.

Periphytic algal communities are highly influenced by substrate types in small pond, not large floodplain lake

Periphytic algal communities are highly influenced by substrate types in small pond, not large floodplain lake

Periphytic Algal Succession with Temperature and Light Treatments

Periphytic algae are part of algal periphyton that are important in the microorganism community in the aquatic systems as they play an essential role as producers in the trophic and aquatic levels. The aim of this study serves to determine the structural differences of periphytic algae communities in both Cipeles River and Cisudajaya River, Sumedang, West Java. Accumulation of periphytic algae colonisation was performed with a number of temperature and light treatments using a square-shaped stone sized 2 cm x 2.5 cm x 1 cm for 28 days. Experiments were carried out with the temperatures set to 30°C and 33°C, as well as light intensities of 1000 lux and 2000 lux. Water as the medium is taken from the Cipeles River and Cisudajaya River. A number of different commercial hydroponic fertilisers as the growth nutrients for the periphytic algae were then added to the media. Once the experiments were fully performed, the results showed structural differences in the periphytic algae community from both Cipeles River and Cisudajaya River water media, where 25 types of periphytic algae from 10 classes were obtained from Cipeles River while the Cisudajaya River had 30 types of periphytic algae from 9 classes. In addition, an increase of colonisation was found in the periphytic algae samples taken from Cipeles River and Cisudajaya River, where they experienced biomass accumulation on Day 14 and Day 28. The optimal temperature for the periphytic algae colonisation in both samples is 30 °C, with the optimal light ranging betwen 1000 lux and 2000 lux

Spatial variation in taxonomic and functional composition of periphytic algae in lakes of Karelian Isthmus

Variation in species and functional structure of periphytic algae community in relation to chemical, physical, climatic and spatial factors was studied in 55 lakes of Karelian Isthmus. To assess factors driving functional structure, all algal species were divided into eight functional groups that differ in cell/colony morphology and attachment mode to the substrate. Variance partitioning analysis showed that environmental factors (chemical and climatic) prevailed over spatial variables in explaining variation in periphyton species structure. On the contrary, periphyton functional structure depended more on spatial variables than on environmental factors (chemical and physical). This suggests that dispersal processes which are dependent on landscape characteristics of the region and hydrological connectivity along individual river networks play a more significant role in spatial variation of functional composition of periphytic algae.

Strong top-down effects of omnivorous fish and macroinvertebrates on periphytic algae and macrophytes in a tropical reservoir

Top-chain predators play a fundamental role in the functioning of shallow reservoirs. Studies to elucidate the cascade effects produced by fish on macroinvertebrates and periphytic algae are fundamental for understanding the functioning of aquatic ecosystems. We conducted an enclosure experiment to test the isolated and combined effects of macroinvertebrates and Prochilodus brevis (omnivorous fish) on periphytic algae and the floating macrophyte Eichhornia crassipes for 30 days in the Jazigo reservoir, Brazil. The predation of P. brevis on macroinvertebrates increased the biomass of periphytic algae during the experiment, while the effects of isolated macroinvertebrates decreased the biomass of periphytic algae and increased the biomass of E. crassipes. Macroinvertebrates (which freely colonized the enclosures) were reduced by fish; however, the biomass of the periphytic algae was not reduced. Our results showed that P. brevis has strong effects on the structure of macroinvertebrate and periphytic algae communities, with a visible cascade effect on the abundance of macroinvertebrates. This study contributes to our understanding of the ecology of tropical lakes and the regulation of periphytic algae and macrophyte growth.

Algal Community Dynamics and Underlying Driving Factors in Some Crenic Habitats of Kashmir Himalaya

Given the authoritative and well-documented publication records that crenic habitats support the substantial aquatic biodiversity, understanding of algal dynamics in response to anthropogenic and natural stressors in these crenic systems seems paramount. We sampled and monitored twelve freshwater springs for a period of 2 years from 2014 to 2015 to observe algal dynamics and the factors govern the distribution and dynamics. We used ANOVA, nMDS, PCA, ANOSIM, SIMPER, and BIOENV to reveal the key physicochemical variables influencing the distributional pattern and dynamics of algae. The analysis of variance (ANOVA, Tukey’s post hoc test) revealed significant difference among the springs with dominance of Bacillariophyceae (62%) followed by Chlorophyceae (18%) whereas nMDS ordination of abundance data in two-dimensional space resulted in a significant separation between spring sites (stress value of 0.13). One-way nested ANOSIM produced a significant distinction between periphytic algal communities in springs (global test R = 0.928, p = 0.001). The results of SIMPER revealed the highest average dissimilarity (60.95%) between springs S4 and S5, with the top five contributing families including Cyanophyceae (30.25%), Bacillariophyceae (25.98%), Rhodophyceae (16.74%), Chlorophyceae (13.64%), and Chrysophyceae (13.39%). BIOENV analysis of the periphytic algal data suggested that the assemblage pattern in all crenic habitats were controlled by discharge, conductivity, dissolved oxygen, total alkalinity, and total phosphorus. Since, springs are groundwater-dependent ecosystems acting as ecohydrologic refugia, any small change in groundwater discharge could strongly influence the ambient conditions (including water quality and temperature), which in turn influences the biological assemblage patterns and ecosystem services. Therefore, changes in discharge may provide information on possible future ecological change in the springs in relation to rising aridification.

Periphytic Algae Community Structure and Its Relation to Environment Factors in the Main Stream of the Songhua River from 2014 to 2019

Periphytic algae are often used as an indicator to evaluate water quality. Here, the community structure of periphytic algae and its relationship with environment factors were analyzed in the main stream of the Songhua River during the summers of 2014 to 2019. The status and trends in ecological water quality were also evaluated based on bioassessments. Phytoplankton species belonging to 4 phyla and 58 genera were recorded, including 28 Bacillariophyta genera, 17 Chlorophyta genera, 10 Cyanophyta genera, and 3 Euglenophyta genera; Bacillariophyta, Chlorophyta, and Cyanophyta accounted for 48.28%, 29.31%, and 17.24% of the community, respectively. Cell densities varied between 1.29×104 and 8.42×104 ind·cm-3, with an average of 4.35×104 ind·cm-3. The dominant genera were Cyclotella, Melosira, Asterionella, Cymbella, Synedra, Pinnularia, Navicula, and Scenedesmus. The physicochemical water quality showed notable changes during the past six-year monitoring period. Specifically, the dissolved oxygen content increased year on year; ammonia nitrogen, total phosphorus, and total nitrogen first increased and then decreased; and, overall, water quality significantly improved in 2019. Relationship between periphytic algae and environmental factors was further examined using redundancy analysis (RDA), which showed that time was the main factor driving the succession of algal community structure. Dissolved oxygen (DO), ammonia nitrogen (NH4+-N), total nitrogen (TN), total phosphorus (TP), five-day biochemical oxygen demand (BOD), and chemical oxygen demand (COD) were also important environmental variables affecting algal community structure.