Tauchuntersuchungen zur Vertikalverteilung der sublitoralen Helgoländer Algenvegetation

Zusammenfassung 1. Die Vertikalverteilung der makroskopischen Algen im Helgoländer Sublitoral wurde an fünf Tauchprofilen von insgesamt 7200 m Länge untersucht. Weitere Tauchgänge wurden durchgeführt, um auch seltenere Arten zu erfassen.2. Zwischen 0,5 und 1,5 m Tiefe bildenLaminaria digitata undL. saccharina eine geschlossene Vegetation, zwischen 1,5 und 4 mL. hyperborea. Mehrjährige Exemplare vonL. digitata undL. saccharina kommen nicht unterhalb von 3 m Tiefe vor,L. hyperborea wurde bis 8 m Tiefe gefunden.3. Insgesamt wurden von 1965 bis 1970 im Helgoländer Sublitoral 66 Algenarten bzw. nicht weiter identifizierte Gattungen gefunden (Tab. 2). 38 Arten kamen in mehr als acht der untersuchten 1232 Quadrate (25×25 cm) vor (Tiefenverteilung in Abb. 10). Die sublitorale Algenvegetation erstreckt sich bei Helgoland bis 15 m Tiefe. Durch Vergleich mit Daten von anderen Küsten wurden zwei Artengruppen herausgestellt, deren untere Tiefengrenzen einerseits im oberen, andererseits im unteren Drittel des Sublitorals zu finden sind (Tab. 5).

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Diving investigations on the vertical distribution of sublittoral algal vegetation near Helgoland

The vertical distribution of macroscopic sublittoral algae near the island of Helgoland (North Sea) has been investigated using theScuba diving technique.Laminaria digitata andL. saccharina occupy the depth range between 0.5 and 1.5 m (below mean low water of spring tides). TheL. hyperborea forest extends between 1.5 and 4 m depth. The deepest-growing specimens ofL. hyperborea have been recorded at 8 m depth, the deepest algae (encrusting forms) at 15 m. Some species are confined to the upper part of the sublittoral zone (e. g.Chondrus crispus, Dumontia incrassata, Ahnfeltia plicata, Polyides rotundus, Furcellaria fastigiata), while others penetrate to the lower part (e. g.Phycodrys rubens, Delesseria sanguinea, Brongniartella byssoides, Phyllophora crispa, Bryopsis plumosa).

     

Macroalgal species diversity and biomass of subtidal communities of São Miguel (Azores)

The benthic algal communities of two subtidal sites, located on opposite coasts of S?o Miguel Island (S?o Roque in the south and S?o Vicente on the north coast), were studied over a 2 year period (September 1993–September 1995). At both sites the sublittoral region was surveyed from low-tide level down to a depth of 15 m. Qualitative and quantitative changes are described. A depth-related gradient in species diversity and biomass was found at both localities. In general, red algae such as corallines and Pterocladiella capillacea predominated in the shallow sublittoral (5 m) while brown algae such as Zonaria and Stypocaulon were more abundant at 15 m. Multivariate analyses emphasized the existence at each study site of two communities (5 and 15 m depth), separated by a large transition zone. The 15 m community on the south coast site showed the largest number of species (52), whereas the lowest diversity (30 species) also occurred at this site in the 5 m community. A clear seasonal pattern of biomass change could be discerned only at S?o Vicente where the highest biomass was recorded in spring/summer. No major inter-annual variations could be detected, indicating relatively stable communities at least on a short-term basis. Received in revised form: 28 March 2001 Electronic Publication     

Patterns of macroalgal diversity, community composition and long-term changes along the Swedish west coast

This study analyses the complicated patterns of vertical distribution of the macroalgal vegetation in an area where brackish and marine waters meet and mix. Variables used to record vegetation characteristics are algal cover, species composition and diversity. The data set includes 64 diving profiles, all from sites exposed to wave action, along a ca. 260 km long coastline. The profiles belong to four categories: coastal sites in the Skagerrak (more marine), coastal sites in the Kattegat (more brackish), coastal sites in the Kattegat after a toxic phytoplankton bloom, and submerged offshore stone reefs in the Kattegat. The highest species diversity was found at the reefs, which are not affected directly by land runoff. At the reefs the 18 most common perennial species penetrate 2–11 m (on average 5.5 m) deeper than at the coastal sites. The virtual absence of sedimentation, and thus the availability of substratum, at the reefs may explain the differences so that the lower limit for the algae is determined by light penetration or by recruitment problems caused by strong currents at the reefs, whereas sedimentation limits the settlement of algae in coastal sites. Ordination analysis based on species composition reveals that the major environmental gradients structuring the algal vegetation in the Kattegat and the Skagerrak are salinity and water depth. The large data set of this study made it possible to quantify the downward dislocation of Atlantic intertidal species to the sublittoral along the Swedish west coast. For example, the mean upper limit of Corallina officinalis is 2 m in the Skagerrak but 12.5 m in the Kattegat and the mean occurrence interval of Fucus serratus is 0.9–2.7 m in the Skagerrak, but 1.1–6.3 m in the Kattegat. This downward dislocation is suggested to be the result of decreased competition when species successively disappear with lower salinity. Comparisons of the present study's results with those of previous investigations show that eight common red algal species have moved upwards compared to the situation before the large-scale eutrophication started in the 1960s, e.g. Cystoclonium purpureum and Polysiphonia elongata by ca. 8 m, Phycodrys rubens and Delesseria sanguinea by ca. 5 m. A toxic phytoplankton bloom affected macroalgal community composition on the whole only slightly, but it had a negative effect on algal cover and species richness below a water depth of ca. 5 m, the algae were visibly damaged and the lower vegetation limit temporarily moved upwards.     

LINKING BENTHIC ALGAL BIOMASS TO STREAM SUBSTRATUM TOPOGRAPHY1

The physical properties of substrata significantly influence benthic algal development. We explored the relationships among substratum surface texture and orientation with epilithic microphytobenthic biomass accumulation at the whole‐substratum and micrometer scales. Unglazed clay tiles set at three orientations (horizontal, vertical, and 45°), and six substrata of varying surface roughness were deployed in a prairie stream for 3 weeks. Substrata were analyzed for loosely attached, adnate, and total benthic algal biomass as chl a, and confocal laser scanning microscopy was used to measure substrata microtopography (i.e., roughness, microscale slope angles, and three‐dimensional surface area). At the whole‐substratum level, vertical substrata collected significantly (P < 0.05) less algal biomass, averaging 34% and 36% less than horizontal and 45° substrata, respectively. Benthic algal biomass was also significantly less on smoother surfaces; glass averaged 29% less biomass than stream rocks. At the microscale level, benthic algal biomass was the greatest at intermediate values, peaking at a mean roughness of approximately 17 μm, a mean microscale slope of 50°, and a projected/areal surface area ratio of 2:1. The proportion of adnate algae increased with surface roughness (26% and 67% for glass and brick, respectively), suggesting that substratum type changes the efficiency of algal removal by brushing. Individual substrata and microsubstrata characteristics can have a strong effect on benthic algae development and potentially affect reach scale algal variability as mediated by geomorphology.     

Ecology and dynamics of two intertidal algal communities on the littoral of the island of São Miguel (Azores)

The intertidal benthic algal communities of two sites located on opposite coasts of São Miguel Island (Azores), were studied over a 2-year period (September 1993–September 1995). At both sites (São Roque on the south coast and São Vicente on the north), the littoral region was surveyed from the upper intertidal down to the sublittoral fringe. The survey revealed five distinct zones, with a variable degree of overlapping. The two upper zones were characterized by animals (littorinids and barnacles, respectively). Lower down, algal communities formed three distinct zones: an upper Fucus spiralis/Gelidium microdon association, a more extensive turf zone, and a belt featuring erect or frondose algae. Upper in the eulittoral, the turf was mainly monospecific, and dominated by Caulacanthus ustulatus. The lower eulittoral turf was dominated by articulated coralline algae, the associated species differing between the two sites studied. Mainly erect algae (Pterocladiella, Asparagopsis, etc.) occurred furthest down the shore and extended into the nearby shallow sublittoral. Two intertidal communities were studied at each locality: the upper eulittoral (Caulacanthus turf in São Roque and the Fucus spiralis/Gelidium microdon association at São Vicente), and the lower eulittoral (the coralline turf). The lower littoral communities had a higher algal diversity. A general pattern was observed in the seasonal variation of biomass: the lower levels exhibited the higher values in late summer/early autumn, the period in which the upper levels had the lowest standing crops. Physical factors are proposed to account for this. No significant inter-annual variations could be detected, indicating relatively stable communities, at least on a short-term basis.     

The macrofauna and macroflora associated withLaminaria digitata andL. hyperborea at the island of Helgoland (German Bight,North Sea)

This paper describes the macroflora and macrofauna associated with two bull kelp species,Laminaria hyperborea andL. digitata, at the island of Helgoland, North Sea. During a study period of seven months (March–September 1987), 29 macroflora species and 125 macrofauna species were found. The dominant taxonomic groups were Polychaeta (25 species), Bryozoa (17), Amphipoda (14), Hydrozoa (10) and Ascidiae (8). The species maximum was in July. In general,L. hyperborea was preferred as a substrate for settlement toL. digitata. Composition of the communities associated with kelp changed during the season according to exposure to wave action, and according to location on the kelp thallus. The rhizoid community of both kelps bore more species at exposed locations. Wave-exposedL. digitata lacked obvious faunal settlement on both phylloid and cauloid. Phylloid and cauloid ofL. hyperborea were chosen as an attractive substrate at both sheltered and wave-exposed locations, showing an association of encrusting bryozoan and hydrozoan colonies.     

A subtidal transect in Jervis Bay,New South Wales

Long-term variations in the intertidal algal flora of the entire rock platform at Plantation Point, Jervis Bay, are described by May (1981). The study now reported presents similar long-term changes in the flora of a subtidal region of the same headland, observed within the same period of time. This first detailed report of subtidal macroalgal communities in New South Wales describes a several-year study of the benthic communities along a transect in the upper sublittoral region of a rocky headland at Plantation Point, Jervis Bay. Eighty-nine species of algae were recorded, five of which were previously unrecorded for New South Wales. The area studied is dominated by the large brown algae Ecklonia radiata and Phyllospora comosa, large areas of which were cleared periodically by storms. Turf, shade and crustose coralline algal communities also were present. Storms, seasonal variation and longer term changes all affected the abundance and distribution of the algal species growing along the transect and hence the floristic composition of the area.     

Bacteria biofilm encourages algal immigration onto substrata in lotic systems

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

扎龙湿地藻类优势种生态位及其种间联结性动态分析

为了解扎龙湿地藻类群落结构的季节特征和演替规律,对扎龙湿地藻类群落进行了分析,主要涉及藻类优势度、生态位、MFG功能群划分、生态位重叠值及联结系数分析。结果表明：(1)在检出的8门311种藻类中,夏季种类丰富度最高,全年优势种为狭形纤维藻和啮蚀隐藻。(2)优势种种类和密度季节性差异较大,春季以金藻-绿藻种类为主,夏季以绿藻-隐藻种类为主,秋季以隐藻-绿藻种类为主。(3)生态位宽度值与MFG功能群划分结合分析表明,优势种可分为3个大类别,不同种类对资源利用情况的差异较大,各季节广生态位种能较好的反映扎龙湿地水体环境的季节变化趋势。(4)生态位重叠值在不同季节上有较大差异,O_ik>0.6的占比分别为42.86%(春)、28.57%(夏)和25.00%(秋),表明春季优势种种间资源的竞争最为突出,种间竞争强于夏、秋两季。(5)种间联结性检验结果显示,扎龙湿地藻类优势种间总体联结上呈正关联关系,但是种间联结较为松散,种与种之间相对独立。研究表明,扎龙湿地藻类群落结构生态位测度均存在明显的季节波动,较好反映出水环境季节变化特征,可用来指示湿地水体环境的变化。     

Diel vertical movements and feeding behaviour of blue humphead parrotfish Scarus ovifrons in a temperate reef of Japan

The feeding ecology of scarinine parrotfishes on tropical coral reefs has received considerable attention in the past few decades; nonetheless, relatively few studies have been conducted in high-latitude reefs. Among the Indo-Pacific Scarus species, Scarus ovifrons is unique, being largely restricted to the warm temperate waters of Japan. Nonetheless, there is very little information available on the feeding ecology of this species. In this study, the authors used acoustic telemetry to detect the diel vertical movement patterns of S. ovifrons, video survey to detect its feeding depths and substrata and focal follow survey and genetic analysis to identify algae composition on the feeding scars at Kashiwajima Island, southwestern Japan (32° 46′ N, 132° 38′ E). Acoustic telemetry revealed that S. ovifrons spent most of its time in shallow water (<10 m) during the day and slept in deeper water (10–15 m) at night. Video and focal follow surveys revealed that most fishes of various sizes regularly took bites on epilithic algae and detrital materials on rocky substrata at depths of <10 m, but large fishes (>40 cm total length) sometimes took bites directly on live corals (Acropora solitaryensis) at the 5 m depth zone where live tabular corals dominated the benthos. Molecular phylogenetic analyses revealed that epilithic algae collected from feeding scars were mainly composed of Rhodophyta, and coralline algae were less often targeted. Overall, this study revealed that S. ovifrons feeds mostly at depths <10 m, and the feeding algae substrata of the species are similar to those of tropical coral reef parrotfishes.     

Algal epilithon and water quality of a stream receiving oil refinery effluent

Changes in epilithic algal communities colonizing introduced substrata were determined in a stream polluted with oil refinery effluent at Digboi (Assam, India). The number of algal taxa was reduced but the growth of blue-green algae, particularly two species ofOscillatoria, was encouraged. Epilithic biomass (as chlorophylla) also declined at polluted stations. The algal community of the upstream station was markedly different from the community occurring just after the confluence of effluent; however, the differences were gradually reduced downstream, indicating improvement in water quality. Of the various criteria tested for possible relationships with the level of pollutants, species richness, Shannon diversity and biomass showed significant relationships. The study demonstrates the usefulness of algal criteria for monitoring oil pollution in running waters.     

Grazing effects in macroalgal communities depend on timing of patch colonization

Rocky macroalgal assemblages are typically composed of patches differing in age and species composition and grazing is generally a very important modifier of such assemblages. We hypothesized that patch colonization time determines its algal community and that grazing effects depend on the colonization time and vary with depth. We created patches by placing empty substrates at two sublittoral depths over five consecutive months, manipulated grazer entry and determined the algal species composition in each patch in the next growing season. Distinct algal colonization periods resulted in different algal assemblages. Although algal communities in our study area consist mainly of opportunistic species, thus being highly dynamic, the resulting macroalgal assemblages differed in species richness, diversity, composition, and total biomass even a year after first colonization. Substrates close to the water-surface supported a higher species richness and diversity than those in the deeper littoral. The community characteristics, total density, total biomass and species richness were only slightly, if at all affected by grazing. However, individual algal species or taxa showed varying and even contrasting responses to grazing, often differently between depths and depending on colonization time. In the deeper littoral, but not close to the water surface, grazing increased the density of filamentous brown algae while reducing the green alga Cladophora glomerata. In these taxa, grazing effects were strongest in patches colonized during the early growing season. Grazing at the colonization stage had lasting consequences for the density of several individual species.     

A quantitative scuba-diving survey of the sublittoral macrobenthos at subantarctic Marion Island

Summary A SCUBA-diving survey of the macrobenthos of hard substrata in the sublittoral zone at subantarctic Marion Island was conducted during March and April 1988. Dense beds (12 kg m^–2) of the kelp Macrocystis laevis occur in depths > 5 m. Durvillaea antarctica is found along the infralittoral fringe and Desmarestia rossi and Durvillaea sp. occur in a narrow zone from 3 m–6 m. Under-storey algae (chiefly rhodophytes) tend to decrease in biomass with depth, with mean values of 1.57 kg m^–2 at 5m, 0.75 kg m^–2 at 10m and 0.49 kg m^–2 at 15 m. Encrusting coralline algae are particularly abundant in shallow areas (¯x = 0.92 kg m^–2) but are insignificant in deeper areas. Total biomass of macrozoobenthos increased with depth with mean values of 0.12 kg m^–2 at 5 m, 0.34 kg m^–2 at 10 m and 0.46 kg m^–2 at 15 m. Polychaetes, crustaceans, echinoderms, molluscs, sponges and bryozoans dominated the macrozoobenthos in terms of biomass. Approximately 200 species of macrobenthic animals were recorded and numerically, polychaetes, crustaceans, molluscs, nematodes and echinoderms dominated. The sublittoral benthos at Marion Island is compared with that occurring at other subantarctic and Antarctic islands, in particular, the Kerguelen Island group. Zoogeographic trends and the possible effects of nutrient input from seabird guano are briefly discussed.     

Die Makrofauna und ihre Verteilung im Nordost-Felswatt von Helgoland

The macrofauna and its distribution in the sheltered, rocky intertidal zone of Helgoland (North Sea) was studied at 9 vertically and/or morphologically different stations from March to September in 1984. Seasonal variations in the communities were described based on each species' conspicuousness. A total of 172 species was found. The macrofauna shows a zoned pattern, but also the different substrata, for example, affect its distribution. The number of species increases from the upper intertidal to the upper sublittoral zone from 23 to 133 species. The upper intertidal is characterized byLittorina saxatilis, Chaetogrammarus marinus andHyale nilssonii. Typical and abundant species of the middle and lower intertidal areFlustrellidra hispida, Littorina mariae/obtusata, Littorina littorea, Mytilus edulis andSpirorbis spirorbis. The upper sublittoral zone is characterized byGibbula cineraria and increasing species numbers of Bryozoa, Nemertini and Opisthobranchia. Only few species (e.g.Dynamena pumila, Laomedea flexuosa, Polydora ciliata, Fabricia sabella, Jaera albifrons, Carcinus maenas) occur in the entire intertidal zone. In comparison to other very sheltered shores in Great Britain, which are also dominated by Fucaceae, the macrofauna in the Helgoland intertidal zone lacks several littoral species, such asPatella spp.,Monodonta lineata, Gibbula umbilicalis, Littorina neritoides, Chthamalus spp., whereasLittorina littorea andGibbula cineraria are highly abundant.     

Antarctic patterns of shallow subtidal habitat and inhabitants in Wilke’s Land

Studies of east Antarctic marine assemblages on hard substrata are rare. In relation to sea-ice breakout, we assessed benthic patterns of habitat and inhabitants between islands and bays at each of two depths (6 and 12 m) across the Windmill Islands coast. Island sites experience sea-ice breakout in the austral spring, while bay sites typically retain sea-ice cover into the summer and in some places the cover is virtually permanent. Composition of assemblages differed between sheltered bays and exposed islands. Islands were dominated by macroalgae, which also varied with depth. Immediately below the ice–foot zone at 6 m, substratum space were monopolised by foliose red (Palmaria decipiens) and foliose brown (Desmarestia sp.) algae, whereas at 12 m large canopies of Himantothallus grandifolius was abundant. The understorey consisted of a mixture of turfs and encrusting red algae at 6 m, and coralline algae at 12 m. Sheltered bays had large areas of sediment/algal complex and no canopy-forming macroalgae. We found more sponges and hydroids in bays, and more brittle stars around islands. Experiments testing factors that covary with exposure and depth in Antarctica, such as light, sedimentation and ice scour are necessary to determine processes that maintain these striking patterns.     

Adhesion of algal cells to surfaces

This paper reports the cell–substratum interactions of planktonic (Chlorella vulgaris) and benthic (Botryococcus sudeticus) freshwater green algae with hydrophilic (glass) and hydrophobic (indium tin oxide) substrata to determine the critical parameters controlling the adhesion of algal cells to surfaces. The surface properties of the algae and substrata were quantified by measuring contact angle, electrophoretic mobility, and streaming potential. Using these data, the cell–substratum interactions were modeled using thermodynamic, DLVO, and XDLVO approaches. Finally, the rate of attachment and the strength of adhesion of the algal cells were quantified using a parallel-plate flow chamber. The results indicated that (1) acid–base interactions played a critical role in the adhesion of algae, (2) the hydrophobic alga attached at a higher density and with a higher strength of adhesion on both substrata, and (3) the XDLVO model was the most accurate in predicting the density of cells and their strength of adhesion. These results can be used to select substrata to promote/inhibit the adhesion of algal cells to surfaces.     

Comparison of attached algal communities on natural and artificial substrata along a thermal gradient

Colonization rate and community structure of periphyton assemblages was examined on aluminium and glass substrata and compared to populations on four submerged macrophyte species in three temperature zones in Cholla Lake, Arizona, U.S.A. Higher densities were achieved over shorter incubation intervals in the warmer zones (26–35° C). Representatives from the planktonic diatom community were first to colonize artificial substrata during the initial two hour incubation period in all temperature zones. Two periphyton diatom representatives, Amphora coffeiformis and Cocconeis placentula var. lineata were the numerical dominants after one week. Cocconeis placentula var. lineata was most competitive on natural substrata at temperatures <26°C, while Amphora coffeiformis dominated temperature zones >26°C with no significant preferences for artificial or natural substrata. The significance of temperature, specific conductance and availability of living hosts is discussed with respect to regulating populations of these two common periphytic diatom species in alkaline waters in southwestern U.S.A. Similarity indices (SIMI) were used to compare algal assemblages on various natural and artificial substrata pairs. Periphyton assemblages were very similar on all natural substrata within similar temperature zones, with little or no preference for macrophyte species displaying similar leaf morphology. Diatom assemblages were quite similar on aluminium and glass substrata throughout the incubation period in all temperature zones, while blue-green algal populations were significantly different, particularly in the higher temperature zones (>28°C). Natural periphyton communities were best represented after four weeks incubation with aluminium substrata in warmer temperature zones (>28°C) or where filamentous blue-green algae dominated. The selection of adequate incubation time when employing artificial substrata to evaluate natural assemblages for different environmental conditions and algal populations is discussed.     

Observation of developmental processes in loosely attached diatom (Bacillariophyceae) communities

The succession of attached algal communities was studied in Lake Biwa and the Omiya River. After the ‘climax’ which was characterized by large numbers of Cymbella turgidula Grunow and Gomphoneis quadripunctatum (Østrup) P. Dawson, the community was dominated by species attached directly to stalks or tangled among stalks. Stalks produced by species of the previous phase are important as secondary substrata. The dominant species of the new phase were ‘loosely attached species’, as defined by Hudon and Bourget (1983). I propose a new model consisting of phase 1, characterized by the first colonizers, including large rosette‐type species and motile type species; phase 2, characterized by the stalk‐producing species; and phase 3, characterized by the growth of loosely attached species.     

Periphyton Architecture and Susceptibility to Grazing by Periwinkles (Littorina littorea,Gastropoda)

Periphyton was grown on transparent plastic substrata in the Kiel Fjord and used for short-term laboratory experiments to study the feeding selectivity of the periwinkle Littorina littorea in response to the vertical structure of the periphyton. The susceptibility of algae to periwinkle grazing was assessed by comparing the species-specific biomass within the grazing tracks of the snails to the biomass outside the tracks. After 3 weeks of incubation, the periphyton consisted of a scattered monolayer of algal cells without vertical structure. No apparent grazing could be found. After 6 weeks of incubation, periphyton consisted of a tightly attached undergrowth (mainly Cocconeis scutellum, Bacillariophyceae, and Myrionema sp., Phaeophyceae) and canopy of filamentous (Melosira moniliformis, Bacillariophyceae) and stalked forms (Achnanthes longipes, Bacillariophyceae). The unicellular diatoms Fragilaria tabulata and Stauroneis constricta grew partly on the primary substratum and partly as epiphytes on the canopy species. The canopy species and the epiphytes were decimated inside the grazing tracks, while the tightly attached undergrowth species appeared ungrazed.     

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

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