The impact of small chironomid grazers on epiphytic algal abundance and dispersion

• 1 A combination of field and laboratory experiments was used to assess the impact of chironomid grazers on taxonomic composition, abundance and dispersion of epiphytic algal assemblages.
• 2 In the laboratory, Psectrodadius sp. reduced the biovolume of algal species preferred as food and increased the degree of clumping of non-preferred species. Thienemanniella cf. fusca had both positive and negative effects (depending on the algal species) on the biovolumes of algal species preferred as food and increased the degree of clumping of non-preferred species.
• 3 In field exclosures, no effect of removal of chironomid larvae from the grazer assemblage could be detected in autumn or winter experiments. A third, longer removal experiment, conducted in summer, resulted in increased biovolumes of edible Cosmarium spp. and Aphanocapsa spp., preferred foods of chironomid larvae. Biovolumes of Lyngbya sp., Eulbochaete spp. and Oedogortium spp., filamentous taxa used extensively in larval case construction, also increased. Chironomid larvae had no effect on total algal biovolume or biovolume of large unicellular algae.
• 4 Chironomid larvae can influence epiphytic algal assemblages through selective grazing by reducing the biovolumes of preferred foods and through case-building activity by reducing the biovolumes of construction materials.

     

The effects of temperature on feeding in zooplanktivorous walleye,Stizostedion vitreum,larvae

Synopsis We examined the prey capture process in walleye larvae (9.8–18.0 mm mean length) feeding on zooplankton (density 1501^–1) in laboratory aquaria at 15,18.5 and 22°C. Larvae were starved for 8 h prior to the experiment and only allowed to feed for 10 min during the experiment in order to minimize the influence of gut processing or satiation on feeding behaviour. Prey consumption (g min^–1) increased exponentially and prey capture success (%) increased logarithmically with mean walleye length. Prey consumption and attack rate (strikes min^–1) increased significantly with increasing temperature. The effect of temperature on capture success was slight (2% increase from 15 to 22°C) and not significant. Walleye showed positive electivity for medium-sized prey (0.3–0.6 mm body width, mostlyCeriodaphnia quadrangula), negative electivity for small prey (0.3 mm, mostly cyclopoid copepods) and neutral electivity for large prey (0.6 mm, mostlyDaphnia sp.). Neither prey species electivity nor prey size electivity were significantly affected by temperature. However, prey size electivity did show a fairly large effect size with respect to temperature and we suggest that this relationship should be examined further. These results indicate that temperature affects prey capture in walleye larvae primarily by influencing attack rate.     

Selective feeding of four zooplankton species on natural lake phytoplankton

Grazing experiments evaluated by microscopical counting were conducted with different size classes of Daphnia hyalina, D. galeata, Eudiaptomus gracilis, and Cyclops sp., all from Lake Constance, using natural lake phytoplankton as food. Species-specific grazing selectivity coefficients were calculated for the dominant phytoplankton species from weekly experiments. Specific selectivities were found to be largely invariant through the growing season. All zooplankters grazed more efficiently on phytoflagellates such as Rhodomonas and Cryptomonas than on coccales such as Chlorella and Scenedesmus, regardless of their relative abundance in the phytoplankton assemblage. Filtering rates did not decrease in the presence of filamentous algae. Certain filamentous species were grazed efficiently, but only by D. hyalina: Anabaena planktonica, Oscillatoria amphigranulata, and Stephanodiscus binderanus. Large diatom colonies like Asterionella formosa and Fragilaria crotonensis were grazed well only by Cyclops sp. Some algal species were consistently selected against: Mougeotia thylespora and Dinobryon sp. The species-specific selectivity coefficients can be used as weighting factors to assess the effective food concentration relative to Rhodomonas minuta, a reference species for optimal food.     

The removal of a heated water discharge from a lake and the effect upon an epiphytic algal community

The effects of the removal of a heated water discharge from Lake Wabamun, Alberta (longitudes 114°26 and 114°44W; latitudes 53°30 and 53°35N) upon epiphytic algae has been documented. Due to the timing of the cessation of flow the first year without elevated temperatures was one of transition. Standing crops returned to normal lake levels the following year. Species composition changes occurred, and species such as Epithemia turgida, Amphipleura pellucida, Nitzschia fonticola, Rhopalodia gibba, Rivularia haematites, and Scytonema alatum, all of which were detrimentally affected by the elevated temperatures, returned to the previously affected area, while species such as Oedogonium sp., Spirogyra sp. and Cladophora glomerata, dominant at times in the heated area, diminished in importance. The ability of Cocconeis placentula to grow at low light intensities (15–110 µEinsteins m^–2 s^–1) was again demonstrated.     

Extracellular fibril production by freshwater algae and cyanobacteria

In order to study the ability of freshwater algae and cyanobacteria to form extracellular fibrils, a screening test using ruthenium red (RR) staining was carried out on 28 species. Five of these were examined for growth and production of fibrillar material in culture media of different phosphate (P;) contents. RR-staining and uronic acid determinations at various stages of algal growth were complemented by electron microscopy of the cells and of fibrillar material released into the medium. The lower P_i concentrations enhanced growth of Micrasterias radiata, Eremosphaera sp., and Microcystis aeruginosa, and had little or no effect on growth of a Xanthidium sp. and Scenedesmus quadricauda. Extracellular uronic acid production, which was higher in low P_i medium in M. radiata, M. aeruginosa, and Xanthidium sp., could reach levels of 50 mg/liter or more. Algae with high proportions of RR-positive cells (M. radiata, Eremosphaera sp., Xanthidium sp., and M. aeruginosa) produced high levels of slime-like material and distinct fibrils that were often seen attached to the cell surface and only slowly released into the medium. No such material was found in cultures (or supernatants) of Sc. quadricauda, which also produced relatively low amounts of polyuronic acids. Specific types of filaments, often forming fascicles with rectangular arrays of globular particles were observed by negative staining electron microscopy of some algal cultures. RR-positive material was also observed in the cytoplasm and on the cell walls and surfaces of M. radiata and M. aeruginosa.Offprint requests to: D. J. Kushner.     

Protozoan communities in chalk streams

This study assessed the individual effects of three mayflies (Paraleptophlebia sp., Ephemerella subvaria McDunnough and Epeorus sp.) and one caddisfly (Psilotreta sp.) on periphyton communities associated with clay tiles and leaves. Algal densities were estimated for leaf discs and tiles from experimental chambers (with individual grazers) and control chambers (i.e., no grazers). Scanning electron micrographs (SEM) of leaf discs and tiles also were taken for all mayfly grazing experiments. Densities of algae on leaf discs were two to five times lower than on tiles. Mouthpart morphology influenced how different insects grazed the periphyton community. Paraleptophlebia had typical collector-gatherer mouthparts and had no effect on diatom densities associated with leaves whereas diatom densities on grazed tiles were higher than densities on tiles from control chambers. Epeorus had brusher mouthparts and had little impact on diatom densities regardless of substratum type. The other two grazers had the blade-like mandibles of a scraper. Psilotreta did not reduce the numerical abundance of diatoms on either substratum, but did alter community structure by significantly reducing densities of stalked Gomphonema olivaceum and large species of Navicula and Nitzschia; densities of smaller diatoms (Achnanthes spp) increased. However, E. subvaria reduced densities of most algal species regardless of size on both substrata and also significantly altered community structure. SEMs of substrata grazed by mayflies showed reductions in fungal hyphae on all grazed leaf discs, decreases in filamentous algal forms on grazed tiles, and greatly shortened stalks of G. olivaceum (Paraleptophlebia only). Thus, periphyton communities are different on leaves versus tiles and grazers with different mouthpart morphologies have varying effects on both algal and heterotrophic microbial community structure.     

Interactions of Algae within the Community of <Emphasis Type="Italic">Gracilaria gracilis</Emphasis> (Rhodophyta)

Interactions among the unattached red alga Gracilaria gracilis, the dominant species of an algal community, and associated algal species Chaetomorpha linum, Enteromorpha prolifera f. prolifera, and Polysiphonia sp. were studied during and after an algal bloom. It was shown that during their bloom the associated algae Enteromorpha and Polysiphonia sp. significantly decreased the photosynthetic rate of G. gracilis but did not affect its growth rate. It is suggested that the inhibition of Gracilaria gracilis photosynthesis is connected to the impact of extracellular metabolites excreted by Chaetomorpha linum, Enteromorpha prolifera f. prolifera, and Polysiphonia sp. In laboratory experiments, the photosynthetic rate of the associated species was enhanced in the presence of Gracilaria. However, no significant alterations were observed in the content of chlorophyll a, growth, and the dark respiration rates of associated algae when they were kept together with Gracilaria. It was suggested that allelopathic interactions observed among algal species during the formation of the monospecific Gracilaria community, as well as during algal blooms, are not determinative.     

Phytoplankton diet of Arctodiaptomus salinus (Copepoda, Calanoida) in Lake Shira (Khakasia)

A flow-through method was employed to study the algal part of the diet spectrum of the calanoid Arctodiaptomus salinus in the surface layer of the lake and in the phytoplankton biomass maximum zone (10 m deep). Daily consumption rates of this calanoid differed markedly in these layers, being 1 and 11 g ind ^–1 d^–1, respectively. The cyanobacterial taxa, Lyngbya contorta and Microcystis sp., accounted for more than a half of the ration. Nevertheless, the negative Ivlev electivity coefficient indicated that Arctodiaptomus prefers none of these two taxa. The low measured uptakes suggest that the energy expenditure is compensated by other resources, presumably microzooplankton and detritus.     

Interactions between freshwater bacteria andAnkistrodesmus braunii in batch and continuous culture

Batch and continuous cultures ofAnkistrodesmus braunii were established in an inorganic medium with growth rate limited by P. In batch culture, inoculation of lake water bacterial isolates ofPseudomonas sp. andFlavobacterium sp. showed that thePseudomonas isolate was capable of more rapid growth on algal exudates of lytic products than was theFlavobacterium isolate. When inoculated singly into a continuous culture (D=0.267 day^–1; P level, 2M), theFlavobacterium isolate initially caused a decrease in the population density of the alga, but then steady states for both organisms were obtained. ThePseudomonas isolate under the same conditions caused a rapid washout of the algal culture, and steady-state conditions were never obtained. When thePseudomonas isolate was added to the two-member, steady-state system ofA. braunii andFlavobacterium, the algal population again washed out of the vessel, followed by theFlavobacterium and then thePseudomonas isolate. A transient increase in the P concentration to 200M in the culture vessel caused the low algal population level to increase, followed by increases in the bacterial isolates when the algal population was high enough to supply the required organic carbon source. The system demonstrated that competition for P between the alga and the bacteria can occur, and the results were dependent on the algal and bacterial relative growth rates. The bacterial growth rates were limited initially by organic substrates produced by the alga, and the different bacterial isolates competed for these substrates.     

<Emphasis Type="Italic">δ</Emphasis><Superscript>13</Superscript>C values and crassulacean acid metabolism in <Emphasis Type="Italic">Clusia</Emphasis> species from Panama

The genus Clusia is notable in that it contains arborescent crassulacean acid metabolism (CAM) plants. As part of a study of CAM in Clusia, titratable acidities were measured in 25 species and ¹³C values were measured for 38 species from Panamá, including seven undescribed species, and 11 species from Colombia, Costa Rica and Honduras. CAM was detected in 12 species. Clusia flava, C. rosea and C. uvitana exhibited ¹³C values or diurnal fluctuations in acidity indicative of strong CAM. In C. croatii, C. cylindrica, C. fructiangusta, C. lineata, C. odorata, C. pratensis, C. quadrangula, C. valerioi and C. sp. D diurnal fluctuations in acidity were consistent with weak CAM but the ¹³C values were C₃-like. All of the species that exhibited strong or weak CAM were in the C. flava or C. minor species groups. CAM was not detected in any member of the C. multiflora species group. Strong CAM species were not collected at altitudes above 680 m a.s.l. On the basis of ¹³C values, the expression of CAM was similar in terrestrial, hemi-epiphytic and epiphytic species and did not differ between individuals of the same species that exhibited different life-forms. This study indicates that phylogenetic affiliation may be a predictor of an ability to exhibit CAM in Clusia species from the Panamanian region, and that weak CAM is probably a common photosynthetic option in many Clusia species. ¹³C value is not a particularly good indicator of a potential of Clusia species growing in the field to exhibit CAM because it appears that the contribution in most species of CAM to carbon gain is generally rather small when integrated over the life-time of leaves.     

Development of filamentous green algae in the benthic algal community in a littoral sand-beach zone of Lake Biwa

Temporal changes of biomass and dominant species in benthic algal communities were investigated in a littoral sand-beach zone in the north basin of Lake Biwa from December 1999 to September 2000. Chlorophyll-a amounts of benthic algal communities per unit area of the sandy sediments rapidly increased from late April to June. Increases in biomass of the benthic algal communities are considered to result from the propagation of filamentous green algae Oedogonium sp. and Spirogyra sp. The cell numbers of filamentous green algae and chlorophyll-a amounts of benthic algal communities at depths of 30 and 50cm at a station protected by a breakwater in May were significantly higher than those of a station exposed directly to wave activity. Thus, the biomass accumulation of the benthic algal communities seems to be regulated strongly by wave disturbance. The development of filamentous green algae may contribute to the increase in biomass of the benthic algal community and to the changes in seasonal patterns of biomass in the sand-beach zone of Lake Biwa. We consider that the development of the filamentous green algal community in the littoral zone of Lake Biwa is the result of eutrophication.     

Periphyton flora of some lotic and lentic environments of Hope Bay (Antarctic Peninsula)

A taxonomic study was made of the periphytic algal flora of some freshwater environments at Hope Bay, Antarctic Peninsula (63°27S; 56°59W), during summer 1992–1993. Seven lakes and two pools that are characterized by different trophic levels were studied. The largest, Boeckella Lake, has an important outflow (Prasiola Stream) that was also analyzed. Ninety-seven algal taxa, including specific and infraspecific entities, are recorded. Thirty-one new records for the Antarctic continent are described and illustrated. The flora comprises 44.4% Cyanophyceae, 21.6% Bacillariophyceae, 18.5% Chlorophyceae, 9.3% Tribophyceae, 4.2% Chrysophyceae and 2% Zygophyceae. The epilithic flora of Hope Bay shows a high species richness. Phormidiumfragile and Lyngbya lagerheimii were the most abundant species among the Cyanophyceae. In the algal mats we also found Chrysophyceae (mainly Hydrurus foetidus and Chrysococcus rufescens) and the macroscopic chlorophycean Prasiola crispa. Tribonema australis sp. nov. (Tribonematales, Tribophyceae) is put forward as a new species. Hormidium fluitans (Gay) Heering is renamed Klebshormidium fluitans.     

Comparative microhabitat use of cyprinid larvae and juveniles in a lotic floodplain channel

Synopsis The microhabitat of six cyprinids in a braided side-channel of the Upper Rhône River, France, was studied June to September 1985, using point abundance sampling by electrofishing. Correspondence analysis of the samples-by-species data and Jacob&s electivity index (calculated from the availability and exploitation of eight environmental variables) revealed three groups of species: (1) Leuciscus cephalus and Alburnus alburnus, which used relatively similar microhabitat during both larval and 0 + juvenile development; (2) Rutilus rutilus, Leuciscus leuciscus and Chondrostoma nasus, which used different microhabitats as larvae but whose microhabitat overlapped notably during 0 + juvenile development; (3) and Gobio gobio, whose juvenile microhabitat overlapped only marginally with that of other 0 + juveniles (no data were available on larvae). Overlap in microhabitat use by 0 + juveniles increased notably during a period of reduced discharge, when the amount of available vegetal and ligneous structures decreased; while most species exploited the increased area of shallow waters, juvenile A. alburnus appeared to be the only species that exploited the deeper waters off the channel&s steeper banks. Although not quantified, numerous field observations of predation threat, particularly during the period of reduced discharge (which reduced the amount of cover), suggest that predation risk may influence the microhabitat use of young fishes. Compared with similar species in European lakes, the overlap in microhabitat use of these fluvial cyprinids appears to be more intensive.     

Plant growth regulatory metabolites from novel actinomycetes

Metabolites from 796 isolates of aerobic actinomycetes were screened for plant growth regulatory properties using an algal bioassay. These included 266 isolates ofStreptomyces, 28 unidentified actinomycetes, and 502 isolates of novel actinomycetes represented by 18 genera. Algal growth inhibition of 30% was observed with 60 isolates, 37 of which belonged to the genusStreptomyces. Among other inhibitors were 8 isolates ofActinomadura, 6 ofActinoplanes, 2 each of the generaThermomonospora, Streptoverticillium, andPromicromonospora, and 3 unidentified. Metabolites from 70 isolates promoted algal growth by 20%. These included 13 isolates ofMicromonospora, 11 ofStreptomyces, 6 ofNocardia, 5 ofActinomadura, and 4 each ofRhodococcus andThermomonospora. Sixteen unidentified isolates; 3 isolates ofPromicromonospora; 2 isolates each ofActinoplanes, Streptosporangium, andOerskovia; and 1 of Thermoactinomyces peptonophilus-like organism andSaccharomonospora viridis also promoted the algal growth by 20%. The plant growth inhibitory properties of 9 actinomycetes and the growth promoting properties of 6 were demonstrable during the secondary screening on higher plants using chemicals extracted from the culture broth. The metabolites fromMicromonospora, Nocardia, Rhodococcus, Streptosporangium, andOerskovia isolates were associated with plant growth promotion only; those fromStreptomyces were most frequently involved with the growth inhibition.This is Michigan Agriculture Experiment Station Journal Article No. 12191.     

The radiation of spring snails of the genus Belgrandiella in Austria (Mollusca: Caenogastropoda: Hydrobiidae)

Grain density microautoradiography (MAR) was used to study uptake of dissolved amino acids (DFAA) in microalgal communities from a sandy bay on the west coast of Sweden. A mixture of fifteen ³H-labeled amino acids (final concentration 20–80 nmol l^–1) was added to sediment samples collected from two depths (0.5 and 4 m), on five occasions representing different seasons. On all sampling occasions, the microflora was dominated by diatoms (> 85% of the total biomass). Cyanobacteria occurred in the summer and autumn, but never dominated the biomass. Between 5 and 48% of all counted algal cells showed uptake (1–44% of algal biomass). Uptake was recorded for all majors microalgal groups (diatoms, cyanobacteria and autotrophic flagellates). Uptake was more frequent on the shallow site and the highest proportion of cells showing uptake occurred in May. Although uptake was common among both motile and attached growth forms, on several occasions uptake frequencies were higher for attached cells. As the attached fraction is frequently burried out of the photic zone, and not able to rapidly migrate towards light, this implies that heterotrophic capacity should be valuable. Also at species level, some taxa showed higher uptake frequencies than others, for example populations of motile diatom species that displayed a seasonal behaviour (blooming species), such as Nitzschia cf. dissipata and Cylindrotheca closterium. Other species showing frequent uptake were the coccoid colony-forming cyanobacterium Microcrocis sp., and the filamentous cyanobacterium Phormidium sp.. The results suggest that there is a high potential for DFAA uptake in the microphytobenthic community and that for some growth forms and species this could imply a competitive advantage.     

Fatty Acid Patterns in <Emphasis Type="Italic">Chlamydomonas</Emphasis> sp. as a Marker for Nutritional Regimes and Temperature under Extremely Acidic Conditions

Fatty acid profiles were used to characterize nutritional pathways in Chlamydomonas sp. isolated from an acidic mining lake (pH 2.7). Surprisingly, profiles of Chlamydomonas sp. grown in the lab under photoautotrophic, mixotrophic, and heterotrophic conditions at in situ deep strata lake water temperatures (8°C) were very similar, polyunsaturated fatty acids including -linolenic acid (18:33) and 16:43 along with palmitic acid (16:0) being most abundant. Therefore, heterotrophic growth of Chlamydomonas sp. at low temperatures can result in high concentrations of polyunsaturated fatty acids, as previously only described for some psychrophilic bacteria. By contrast, the cultivation of isolated Chlamydomonas sp. at 20°C, reflecting surface water temperatures, provided fatty acid patterns characteristic of the nutrition strategy applied: the concentration of polyunsaturated fatty acids decreased when the growth pathway changed from photoautotrophic via mixotrophic to heterotrophic. Total fatty acid concentration also diminished in this order. Principal component analysis confirmed the significance of FA profiling to mirror nutritional pathways. Lake-water analysis revealed low concentrations of dissolved organic carbon, mainly consisting of polymeric fulvic acids that are unable to support heterotrophic growth of Chlamydomonas sp. Polymeric fulvic acids present in the deeper strata of the lake turned out to be formed in situ on the basis of organic monomers including reduced sulfur-containing ones, as revealed by thermochemolysis and pyrolysis. Growth of Chlamydomonas sp. in the deep chlorophyll maximum is therefore assumed to mainly result from photosynthesis, despite very low photon densities. Phytol-including metabolites proved to be significant biomarkers to indicate the nutritional pathway of Chlamydomonas sp. , -Dicarboxylic acids—light-induced degradation products of unsaturated fatty acids—appeared to be good indicators of photooxidative alterations to the algal species under study.     

Comparative limnology of Sambhar and Didwana lakes (Rajasthan,NW India)

Two alkaline saline inland lakes of Indian arid region were studied during 1984 and 1985, to assess functioning and interaction of various environmental and biological factors. Changes in physical and chemical variables, planktonic composition, chlorophyll content and phytoplankton primary productivity were examined.Salinity in both lakes fluctuated from almost fresh water (1.80), to hypersaline (300) and acted as the main controlling factor for almost all the biotic parameters. Maximum total alkalinities were 2162 mg l^–1 and 2090 mg l^–1, respectively in Sambhar and Didwana lakes. Dissolved oxygen ranged from completely anoxic conditions to maxima of 11.68 and 7.29 mg 1^–1, respectively in Sambhar and Didwana lakes. Nutrient enrichment in the lakes was low.The phytoplankton species composition of Sambhar lake was reduced from an earlier reported 20 genera to only 11 (Nostoc, Microcystis, Spirulina, Aphanocapsa, Oscillatoria, Merismopedia, Nitzschia, Navicula, Synedra, Cosmarium and Closterium). Phytoplankton of Didwana was composed of only 9 genera including Anabaena and Nodularia. Sambhar lake, which once contained Artemia, is now totally devoid of them. On the other hand, Artemia was the most dominant zooplankter in Didwana lake at a salinity range of 15–288. Other zooplankters such as Moina, Cyclops and Brachionus flourished at lower salinity levels in Didwana lake. The seasonal quantitative and qualitative phyto- and zooplankton changes in relation to salinity are documented.     

The role of endosymbiotic algae in photoaccumulation of green Paramecium bursaria

The endosymbiotic unit of Paramecium bursaria with Chlorella sp. photoaccumulates in white, blue-green, and red light (<700 nm), whereas alga-free Paramecia never do. The intensity of photoaccumulation depends on both the light fluence rate and the size of the symbiotic algal population. Photoaccumulation can be stopped completely with 3-(3,4-dichlorophenyl)-1,1-dimethylurea (DCMU), an inhibitor of photosynthetic electron transport. Hence the photosynthetic pigments of the algae act as receptors of the light stimulus for photomovement and a close connection must exist between photosynthesis of the algae and ciliary beating of the Paramecium.     

Detection of inconspicuous epiphytic algae supporting food webs in seagrass meadows

Summary Detritus from common seagrasses and other marine angiosperms may often be a less important basis for estuarine food webs than previously believed. In NW Gulf of Mexico seagrass meadows, epiphytic algae have high productivities, palatability, and a more important trophic role than common large plants have. Interdisciplinary field experiments show (1) intensive night-time ingestion of epiphytes by various invertebrate detritivores, (2) very high productivity of epiphytic algae on seagrasses, and (3) assimilation of epiphytes rather than seagrasses, as measured by ¹³C comparisons. These combined data show that many naturally concentrated and potentially competing invertebrates in Gulf of Mexico seagrass meadows feed largely on the algal overgrowth on seagrass blades, even when such algae appear to be sparse. Primary productivity of these epiphytic algae can equal that of the seagrasses, per blade or per unit biomass. Animal ¹³C values tracked epiphytic values rather than seagrass values when comparisons were made over six sites. These measurements reinforce the view that epiphytic algae can be the primary basis of the food web in seagrass meadows.Contribution No. 608 of The University of Texas Marine Science Institute     

Seasonal changes in an intertidal annual algal assemblage in northern Japan: The role of pre-emption and grazing on algal replacement

The pattern and process of seasonal changes in an intertidal annual algal assemblage were examined at Hiura, northern Japan. Short-term field experiments (<2months duration) were set up to quantify the effects of both grazing and pre-emption on species replacement in the assemblage in three different seasons. An 8-month field experiment was set up to quantify long-term effects, including the indirect effects of both grazing and competitive dominance on the community structure. Results suggested that seasonal change in the algal assemblage resulted from the interaction of abiotic environmental change, competition and grazing. The relative contribution of these factors varied within a short period, presumably as a result of seasonal changes in physical environmental stress, free space availability and grazing pressure. From February to March, when grazer density was low and there was much free space available for algae, the dominant species shifted from foliose green alga Monostroma angicava to filamentous red alga Bangia atropurpurea, because B.atropurpurea grew faster than M.angicava. This species replacement was not influenced strongly by biological interaction but by temporal changes in abiotic environmental conditions. From April to mid May, when there was less free space available for algae in the natural community, the dominant B.atropurpurea decreased with increasing foliose red alga Porphyra yezoensis, because only P.yezoensis was able to invade an area pre-empted by algae. Grazing did not affect this species replacement. After mid May, the two dominant species, P.yezoensis and B.atropurpurea, decreased. Their decline was mainly caused by desiccation stress and was partially affected by grazing.