Preliminary results of a classification of fifty-one selected northeastern Wisconsin Lakes (USA) using indicator diatom species

Diatom species lists were generated for 51 lakes in northeastern Wisconsin, and then classified by two-way indicator species analysis (TWINSPAN). The lakes were initially divided into two main groupings: Group I lakes which were alkaline lakes of moderate to high productivity, and Group II lakes which were acid lakes of low productivity. Group I lakes were further divided into two subgroupings, and four levels. In total, twelve indicator diatom species were recognized during the TWINSPAN lake classification, these were: Achnanthes exigua Grun., A. lanceolata var. dubia Grun., Amphora ovalis var. Affinis (Kütz.) V.H. ex Det., A. perpusilla (Grun.) Grun., Cymbella cistula (Ehrenb.) Kirch., Diploneis elliptica (Kütz.) Cl., Eunotia pectinalis (O.F. Mull.) Rabh., Fragilaria vaucheriae (Kütz.) Peters., Frustulia rhomboides var. crassinervia (Bréb. ex W. Sm.) Ross, Navicula capitata Ehrenb., N. decussis Østr., and N. scutelloides W.Sm. ex Gregory.     

BENTHIC MARINE DIATOM ASSOCIATIONS: UPPER FLORIDA BAY (FLORIDA) AND ASSOCIATED SOUNDS1

Models of the diatom associations found in upper Florida Bay and adjoining sounds have been constructed utilizing Q modal factor-vector analysis and ecologic diversity indices (Shannon index, number of species, evenness). Four distinct associations were defined using Q-mode factor-vector analysis. Two associations were epiphytic, occurring on Thalassia testudinum König—Association I was characterized by Cocconeis placentula Ehr., Association III by Cylindrotheca closterium (Ehr.) Reim. & Lewin and Cocconeis placentula Ehr. The other two associations were epipelic, occurring on the carbonate mud substratum—Association II was characterized by Cyclotella striata (Kütz.) Grun., Rhopalodia gibberula (Ehr.) O. Müller and Surirella fastuosa (Ehr.) Kütz., Association IV by Fragilaria crotonensis Kitton and Cyclotella striata (Kütz.) Grun. The majority of the 161 species identified were present in both the epiphytic and epipelic assemblages. Only 33 species were restricted to the epiphyton, and 18 species restricted to the epipelon. The epipelic assemblage was significantly more diverse than was the epiphytic assemblage. A general trend of increased diversity away from terrestrial environs toward more open areas of water in both the epipelon and epiphyton was also found.     

Succession of phytoplankton in a deep stratifying lake: Mondsee,Austria

Phytoplankton numbers, biovolume, chlorophyll-a and various physico-chemical characteristics were followed at weekly intervals in Mondsee, Austria during the year 1982. Secchi-disk transparency varied from 10 m in winter to 2 m in September. Prior to the onset of stratification phosphate-phosphorus concentration was 4 µg 1^–1 decreasing to undetectable values thereafter. Nitrate-nitrogen dropped from 590 µg 1^–1 to about 100 µg 1^–1 during the same time. The vernal bloom was dominated by Asterionella formosa Hass. which abruptly declined after silicon depletion. Spring growth ceased in early June, when Tabellaria flocculosa (Lyngb.) Kütz var. asterionelloides Grun. dominated. Oscillatoria rubescens D.C. and Microcystis aeruginosa Kütz. dominated summer and early autumn followed by the chrysophyte Dinobryon divergens Imh. and D. sociale Ehr. which formed up to 69% of total biovolume in October. Thereafter diatoms and Cryptophyceae (Rhodomonas lacustris Pascher and Ruttner, Cryptomonas pusilla Bach.) became abundant again.Maximum chlorophyll-a concentration in the epilimnion (16 µg 1^–1) was reached during spring growth of the diatoms. During summer higher chlorophyll-a levels were always associated with the metalimnetic layer of Oscillatoria.Compared with earlier studies, both the total biovolume and the share of Oscillatoria rubescens significantly decreased because of reduced nutrient loading of the lake and wash-out of Oscillatoria (theor. renewal time of the lake: 1.7 years).     

Regulation of phytoplankton dynamics in a Laurentian Great Lakes estuary

The composition and dynamics of phytoplankton populations were examined in Old Woman Creek estuary, Lake Erie (USA). The centric bacillariophytes,Cyclotella atomus Hust.,Cyclotella meneghiniana Kütz., andAulacoseira alpigena (Grun.) Krammer, and the cryptophytes,Cryptomonas erosa Ehren. andRhodomonas minuta var. nannoplanctonica Skuja, dominated the phytoplankton most of the year. Chlorophytes, euglenophytes, and cyanophytes were observed less frequently. Estuarine and Lake Erie phytoplankton were considered distinct populations; lake taxa were largely confined to the estuary mouth and present only in low biomass. Maxima and minima of estuarine phytoplankton coincided with meteorological and hydrological forcing in the form of rainfall and subsequent storm-water inflows, respectively. Distinct population dynamics between the upper and lower estuary following storm events were attributed to the presence/absence of refugia serving as a source for repopulation by opportunistic taxa, fluctuating light conditions in the water column resulting from influx of particulate matter and resuspension of bottom sediments, and nutrient inputs associated with surface runoff and sub-surface interflow. Additionally, agricultural herbicides introduced by storm-water inflows potentially may affect and/or control the growth and physiological responses of individual taxa.     

Beiträge zur Cytologie der Blaualgen

Zusammenfassung Mit Hilfe von Ultradünnschnitten wurde die Substruktur von Phormidium frigidum Fritsch, Phormidium retzii Gom., Oscillatoria limosa Ag., Anabaena variabilis Kütz und Cylindrospermum licheniforme Kütz untersucht.Das Chromatoplasma besteht aus submikroskopischen Lamellen, die bei den einzelnen Arten charakteristische Lamellensysteme bilden. Größere Unterschiede in der Ausbildung und Anordnung der Lamellen bestehen zwischen den Vertretern der Oscillatoriaceen und der Nostocaceen.Das Centroplasma läßt auch im submikroskopischen Bereich keine Abgrenzung gegen das Chromatoplasma erkennen. Es ist dadurch gekennzeichnet, daß das Plasma hier kompakter gelagert ist und die für das Chromatoplasma typischen Lamellen fehlen. Das Grundplasma ist körnig und läßt gewisse kettenförmige Strukturen erkennen.     

Seasonal changes in composition and biomass of epiphytic algae on the macrophyte Ranunculus penicillatus in a chalk stream,with estimates of production,and observations on the epiphytes of Cladophora glomerata

The epiphytic algae on surfaces of the macrophyte Ranunculus penicillatus (Dumort.) Bab. var. calcareus (R. W. Butcher) C. D. K. Cook and on Cladophora glomerata (L.) Kütz growing in the River Itchen at Otterbourne near Southampton were studied between February 1984 and June 1985. The river at this site has a mean flow rate of 0.33 m s^–1, and is about 16 m wide and on average 20 cm deep, with a discharge ranging through the year between 0.34 and 2.46 m³ s^–1. The pH of the river varies little around 8.2, with a mean alkalinity of 236 mg HCO _inf3 ^sup1 l^–1, because of its origin from chalk springs. Ranunculus grows throughout the year, with peaks of biomass in spring and autumn. It forms a very large surface for attachment of epiphytes, and covers on average 40% of the stream bed. Numbers of epiphyte cells removed from Ranunculus ranged through the year between 52 × 10³ and 271 × 10³ cells mm^–2 stream floor, with maximum numbers in April, and a secondary peak in October. This pattern partly reflects fluctuations in the biomass of Ranunculus; the number of cells per unit area of plant surface showed a broader spring peak and lower fluctuations in other seasons. Diatoms formed 65 to 98% of these epiphyte cells, with chlorophytes reaching their peak (10%) in summer and cyanophytes (25%) in autumn. Estimates of biomass of these epiphytic forms, derived from measurements of chlorophyll c, indicate a range between 30 and 100 g dry weight m^–2 of weed bed. Colonisation studies showed that the algae grow and reproduce throughout the year, with a mean generation time of about 5 days, suggesting an annual production of about 3 kg dry weight m^–2 of weed bed, which makes epiphytic algae the principal primary producers in the stream. The numbers and biomass of epiphytic algae on Cladophora are considerably less.The species of epiphytic algae found on Ranunculus were generally similar to those growing on Cladophora, and to epilithic algae on pebbles of the stream bed, but different species were dominant on the different substrata. Algal cells in the water column were all derived from benthic habitats, although their relative abundance was very different.     

Seasonality of phytoplankton in some South Indian lakes

The landscape of South India is dotted with innumerable man-made lakes. They differ vastly in age, physiography, water flow characteristics, chemistry and trophic state, yet maintain a phytoplankton overwhelmingly dominated (43–93%) by blue-green algae; the subdominants are diatoms and/or Chlorococcales and euglenoids. The blue-greens apparently reach them from soils which are known to harbour a rich blue-green flora and several species in common with limnoplankton.South Indian lakes resemble some tropical counterparts in sustaining dense phytoplankton populations all the year round and temperate dimictic ones in showing two annual growth peaks that usually occur in summer (February–May) and the post-monsoon period (October–November), in synchrony with rise in temperature. In the chemically more oligotrophic lakes, the peaks are constituted by Raphidiopsis mediterranea Skuja, Navicula cryptocephala Kütz., Melosira granulata (Ehr.) Ralfs, and others and in hypereutrophic lakes by Microcystis aeruginosa Kütz., Synechocystis aquatilis Sauv., Oscillatoria spp., Burkillia coronata West & West and Euglena acus Ehr. The bimodal seasonality in abundance of phytoplankton reflects in chlorophyll and biomass concentrations although these are not in strict synchrony with each other. At the maxima chlorophyll a and over-dry biomass may rise to 8.5 mg l^–1 and 204 mg l^–1 respectively in highly productive waters. The highest rate of carbon assimilation recorded in such phases is 10.6 g C m^–3 d^–1.     

COMMUNITY STRUCTURE DYNAMICS OF EPILITHIC AND EPIPHYTIC DIATOMS IN A MISSISSIPPI STREAM1

Epilithic and epiphytic diatoms were collected monthly from 18 June 1979 through 19 May 1980 from both shaded and unshaded sites of Sessums Creek, a shallow, oligotrophic and strongly alkaline stream in northeastern Mississippi. The dominant macroalgae were filamentous green algae belonging to the non-mucilage producing genera Cladophora and Rhizoclonium. Ninety taxa were identified with Navicula minima Grun. and Nitzschia dissipata (Kütz.) Grun. accounting for 50% of all valves counted. Other important taxa were Navicula menisculus var. upsaliensis (Grun.) Grun., N. cryptocephala var. veneta (Kütz.) Rabh., Achnanthes minutissima Kütz. and Cymbella turgida (Greg.) Cl. With few exceptions, the more dominant taxa were equally abundant in the shaded and unshaded sites and also in the epilithon and epiphyton. Species diversity (H') and the number of taxa in a sample in all four habitats showed an identical pattern, being lowest in winter and highest in fall. No one pair of habitats was more structurally similar than any-other pair when compared by a similarity index (SIMI). Apparently, differences in light intensity and substrate type had little effect on diatom distribution in Sessums Creek and it was concluded that the locality sampled supported a single, nearly homogeneous diatom community.     

Morphological variability of the diatom Cyclotella atomus Hustedt var. atomus and C. atomus var. gracilis var. nov.

Cyclotella atomus collected from different rivers, reservoirs and lakes shows considerable morphological variability. On the basis of electronmicroscopic (EM) study, we propose Cyclotella atomus var. gracilis Genkal et Kiss as a new variety of this species. Cyclotella atomus var. atomus has short and long radial striae, which are irregularly arranged on the same valve, and the alveolar chambers are not clearly divided from the central, hyaline part of the valve face. In addition to the described basic form, several specimens with other features have also been found and are thought to belong to an undescribed variety. The striae of the new variety are equal in length, and the alveolar chambers are clearly divided from the hyaline central area of the valve face by a curved structure.There are EM micrographs on Cyclotella atomus var. gracilis under different names in different papers. The new variety has several characteristics also found in other species, e.g. Cyclotella caspia Grun., or C. meneghiniana Kütz.     

Seasonal variations of Microcystis populations in sediments of Lake Biwa, Japan

Seasonal variations of colony numbers of Microcystis aeruginosa(Kütz.) Kütz. and M. wesenbergii(Komárek) Komárek in N. V. Kondrat. in sediments of Lake Biwa were investigated over a period of 1 year. At two stations located in the shallow South Basin of Lake Biwa (ca. 4 m water depth), the colony number of Microcystisfluctuated seasonally. The number had a tendency to gradually decrease from winter to early summer, while it increased through mid-summer and autumn. Since the Microcystispopulation in sediment was rather small, intensive growth and accumulation in the water column should be important for the formation of Microcystisblooms in Lake Biwa. Microcystiscolonies in the sediment samples after June were observed to be floating in a counting chamber under a microscope. The observation suggests that the recruitment of Microcystis colonies into the water column mostly occurs in early summer. The number of Microcystiscolonies in the deep North Basin of Lake Biwa (70 – 90 m water depth) was larger than in the South Basin. Because the seasonal variation of colony numbers was not observed in the North Basin, and Microcystiscells do not have gas vesicles, these colonies will not return into the water column. The colonies isolated from the sediment of the North Basin were able to grow in cultured conditions, in the same way as those from the sediment of the South Basin. Therefore, Microcystiscolonies may survive for a long time under stable conditions of low temperature (ca. 8 °C) and darkness, in the sediment of the deep North Basin, accumulating gradually each year.     

Phycoperiphyton in selected reaches of the Upper Mississippi River: community composition,architecture, and productivity

Attached algal communities were studied during ice-free periods along the borders of the main channel and in backwaters of the Upper Mississippi River. Diatoms strongly dominated the phycoperiphyton except during late June through August when the green alga Stigeoclonium was abundant. Two distinct assemblages were apparent: a diverse, complex assemblage during spring and late fall annd a less complex, adnate, two-dimensional summer assemblage dominated by Cocconeis placentula var. euglypta and Navicula tripunctata var. schizonemoides. Simultaneous studies revealed two-dimensional periphytic colonization in the main channel and more complex communities in backwaters. Greater physical turbulence (e.g., surface waves) in the main channel may have favored adnate taxa and two-dimensional architecture, whereas lesser turbulence (e.g. protected backwaters) favored the more diverse, complex community. Community architecture and species composition were similar among communities on artificial substrates within each navigation pool (median SIMI0.87), but communities on artificial substrates were not taxonomically very similar (median SIMI=0.44) to epiphytic communities on Cladophora.Accrual of algal cells, chlorophyll a, and aufwuchs ash-free dry weight was usually greater in Pool 5 than in Pool 9. This may have been due to differences in discharges and/or sampler placement. The greatest accrual of cells and chlorophyll a occurred during summer and early fall. Chironomid and tricopteran larvae were common at that time and are known to affect algal accumulation by grazing.     

Seasonal periodicity of planktonic desmids in oligotrophic lake Grane Langsø, Denmark

60 desmids were identified from plankton samples collected during 20 years, bimonthly, or monthly in 5 of the years. Only 5 desmids were perennial, performing cell divisions in a frozen lake at PAR at only 5 cal cm ^–2 day ^–1, below ice-cover. The greatest number of desmids per 100 ml lake water was found in September; it never exceeded 150. Owing to a concentration of free CO₂ of only 0.02 mM l^–1 and a HCO _inf3 ^sup– concentration of 0.002 mM l^–1 the densities of desmid cells were often smaller than 1 cell per 100 ml lake water. A density of 1 cell per 1 ml was attained only by Staurastrum longipes, a desmid showing temporal variation. The relation of the compound phytoplankton quotient to two components of the CO₂-system is discussed. Three new taxa are described: Staurastrum brachiatum Ralfs var. bicorne n. var., Staurastrum thomassonii n.sp. and Xanthidium antilopaeum Kütz. f. bimaculatum n.f..     

An extracellular polysaccharide produced by Palmella mucosa Kütz.

Summary An extracellular polysaccharide composed of glucose, fucose, arabinose and glucuronic acid in a molar proportion of 11:6:3:1 is a major end-product of photosynthesis by Palmella mucosa Kütz.The liberation of polysaccharide is related to the age of the culture. Glucose can substitute efficiently for CO₂ as the source of carbon for polysaccharide synthesis. Nitrate-nitrogen from sodium, potassium and calcium salts can be used in the mineral salts medium with little differences in carbon metabolism of the alga. Ammonium nitrate produces an acidic medium which limits polysaccharide production.The incorporation of C¹⁴ into the polysaccharide from NaHC¹⁴O₃ shows initially a trend toward intracellular synthesis. The C¹⁴ appears in the extracellular polysaccharide after prolonged exposure. Glucose-C¹⁴ is actively transformed to polysaccharide material which is an indication that glucose may play an important role in the synthesis of polysaccharide by Palmella mucosa Kütz.     

Hydrological controls on the diatom assemblage of a seasonal arctic river: Boothia Peninsula,Nunavut, Canada

This study characterizes the melt season diatom assemblage of a middle arctic river with respect to hydrological conditions. In addition, the potential to identify species that show a strong affinity for the lotic environment provides an opportunity to interpret stratigraphic changes in these species in the lake sedimentary record in terms of past hydrological change. Understanding long-term hydrological variability is critical for assessing both current and future environmental change. Significantly higher relative abundances of Achnanthes minutissimaKützing, Fragilaria capucina var. vaucheriae (Kützing) Lange-Bertalot, Diatoma tenuisAgardh, Cymbella arctica(Lagerstedt) Schmidt, C. minutaHilse ex. Rabenhorst, C. silesiaca Bleisch and Encyonema fogedii Krammer in the lotic environment throughout the 2001 growing season compared to the lacustrine sedimentary record suggest that these species characterize the Lord Lindsay River diatom assemblage. Comparison of seasonal abundances of these taxa to hydrological parameters including discharge, electrical conductivity, and water temperature reveal key information about the character of this community. The fact that the river diatom assemblage changes very little throughout the sampling period, despite major changes in hydrological conditions, suggests a degree of resilience and inherent structure in the community. However, a decrease in diatom biomass in response to rapid and dramatic changes in hydrological conditions following a major rainfall event suggests that a threshold tolerance may exist, with potentially important implications for interpreting stratigraphic changes in the paleoenvironmental record.     

A new variety of Nitzschia sinuata (W. Sm.) Grun. from North Wales,Great Britain

Summary A new taxon, Nitzschia sinuata, var. denticuloides, var. n. was found to occur sparingly in two freshwater springs of Newborough Warren, Anglesey, North Wales, Great Britain. These two springs were characterized by shallow waters, moderate temperature and high pH. Dysharmonious or idiotrophic nature of the water was evident from high calcium, sodium and chloride contents, as well as low phosphate and potassium contents. The algal community was found to be dominated by Cladophora fracta (Dillw.) Kuetz. bearing a vast population of epiphytic diatoms such as Epithemia and Achnanthes species.

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Zusammenfassung Eine neue Kieselalge, Nitzschia sinuata, var. denticuloides, var. n., ist in zwei Süsswasserquellen in Newborough Warren, Anglesey, Nord Wales, Grossbritannien spärlich verbreitet entdeckt worden. Diese beiden Quellen sind durch seichtes Wasser, milde Temperaturen und hohen pH Wert gekennzeichnet. Hohe Kalcium, Sodium und Chlorid Werte und niedrige Phosphat und Kalium Werte beweisen den dysharmonischen oder idiotrophen Charakter des Wassers. Die Algengesellschaft bestand hauptsächlich aus Cladophora fracta (Dillw.) Kuetz., dicht besetzt von epiphytischen Diatomeen wie Epithemia and Achnanthes Arten.

     

Interaction of respiratory and photosynthetic electron transport in Anabaena variabilis Kütz.

Prelabeled Anabaena variabilis Kütz. evolves ¹⁴CO₂ in the light with KCN and DCMU (2,4-dichlorophenyl-1,1-dimethylurea) present, comparable to the dark control without inhibitors added. Double-reciprocal plots of CO₂ release vs. light intensity with either KCN or KCN+DCMU present result in two straight lines intersecting at the ordinate. Apparently, reducing equivalents originating from carbohydrate catabolism are channeled into the photosynthetic electron-transport chain, competing for electrons from photosystem II. Under these conditions, the CO₂ release is accompanied by a light-dependent oxygen uptake, presumably due to oxygen-reducing photosystem-I activity while ribulose-bisphosphate carboxylase is inhibited by KCN.Comparing nine blue-green algae it was shown that only nitrogen-fixing species release substantial amounts of CO₂ in the light with KCN or KCN+DCMU present. This release is particularly obvious with Anabaena variabilis Kütz. under nitrogen-fixing conditions, but small when the alga is grown with combined nitrogen.We conclude that nitrogen-fixing species share a common link between respiratory and photosynthetic electron transport. The physiological role may be electron supply of nitrogenase by photosystem I.     

Introduction of a nitrogen-fixing cyanobacterium into tobacco shoot regenerates

Tobacco (Nicotiana tabacum L.) shoots associated with the nitrogen-fixing cyanobacterium Anabaena variabilis Kütz. (ATCC 29413) were regenerated in mixed cultures of tobacco callus and the cyanobacterium. The cyanobacteria were localized inside the tissues as well as on the surface of regenerated shoots, formed heterocysts, and were capable of acetylene reduction.     

A NUTRITIONAL ANALYSIS OF A SUBLITTORAL DIATOM ASSEMBLAGE EPIPHYTIC ON ENTEROMORPHA FROM A LONG ISLAND SALT MARSH1,2

The diatom population structure of a salt marsh epiphytic community growing on Enteromorpha intestinalis was studied at one station throughout the summer. A total of 218 species or varieties were recognized. Six species— Fragilaria construens, Cocconeis scutellum, Cocconeis placentula, Achnanthes hauckiana varieties, Achnanthes pinnata, and Amphora coffeaeformis (var. acutiuscula)–dominated the Enteromorpha epiphytic community during the summer months and comprised ∼ 40% of the total populations. Melosira nummuloides, Opephora martyi, Synedra fasiculata var. tabulata, S. affinis, Navicula platyventris, and N. pavillardi were also very common species (13% of the total population). The distribution of many species in the community was seasonal. A series of differential media have been developed which are effective as tools for the isolation and nutritional characterization of the algae and bacteria from the community. Many diatom species can be recognized by their colony type or growth pattern on solidified media. A key and illustrated plates aid in identification. Changes in population structure of the community were reflected by changes in the nutritional patterns as judged by differences in the growth of diatoms and bacteria on the differential media tested. The nutritional requirements, selectivity, and rank order of media for individual species is given. Almost half of the colony types (32–33) grew on either unenriched seawater from the collection site or basal synthetic seawater. Erdschreiber was a poorer medium for the isolation of algae than seawater alone. Few algal colonies grew on either marine nutrient agar or trypticase soy agar. Nitrite inhibited algal growth. Media enriched with thiamine, biotin, or nitrate, phosphate, and B₁₂ were stimulatory. Soil extract or an acetone extract of Enteromorpha tripled growth; a complex vitamin mixture, or glycerol, or mannitol, or an autoclaved extract of Enteromorpha doubled colony numbers. The greatest numbers of diatom types (43) and total colonies (655) were recovered on media enriched with acetone extract of Enteromorpha, suggesting a possible nutritional relationship between Enteromorpha and its epiphytes. The representativeness of the isolations from the natural community as a function of time was also considered. Concentrations of selected dissolved potential growth-stimulating nutrients were measured within and external to Enteromorpha communities. We conclude that the algal assemblage growing epiphytically on Enteromorpha intestinalis has a diverse auxotrophic profile which contributes to the productivity and stability within this important component of the epiphytic community and that much of the organic substrates used by individual species originate within and are recycled among the community members.     

The epiphytic algae on floating macrophytes of a Paraná river floodplain lake

Summary We describe the composition and seasonal (1993–1994) changes of epiphytic algae on the submerged parts of macrophytes (Eichhornia crassipes (Mart.) Solms-Loubach, Salvinia rotundifolia Willd., Lemna gibba L. and Ricciocarpos natans (L.) Corda) of a shallow lake from Entre Rios province, Argentina (33°48S, 58°57W), permanently connected to the Paraná river.In total, 136 algal taxa were identified: 57 Bacillariophyceae, 31 Chlorophyta, 31 Cyanophyceae, 13 Tribophyceae and 4 Euglenophyceae, all of them characterized by wide geographic distributions. Infertile filaments of Oedogonium spp., Zygnema spp., Spirogyra spp. and Mougeotia spp. (Chlorophyta) dominated the assemblages throughout the year. The diatoms Eunotia bilunaris var. bilunaris, E. monodon var. monodon and two varieties of Gomphonema augur reached high relative frequencies. Characiopsis ettliana and Ch. falx var. cecoalensis (Tribophyceae) were consistently present in low numbers.Multivariate analyses did not reveal major compositional dissimilarities between the four substrates surveyed. Seasonal changes in the community composition were chiefly due to the presence of thychoepiphytic species in association with the interchange of river and lake waters. It is concluded that the pattern of seasonal succession is governed by a complex interaction between hydrometric and seasonal changes.