Interannual variability in the spring phytoplankton bloom in Auke Bay,Alaska

Data on phytoplankton primary production, biomass, and species composition were collected during a 5 yr (1985–1989) study of Auke Bay, Alaska. The data were used to examine the interannual differences in the timing, duration, and magnitude of the spring phytoplankton blooms during each year and to relate these differences to interannual variations in weather patterns. Within any given year, a pre-bloom phase was characterized by low available light, low rates of primary production, low biomass, and predominantly small (<10µm) diatoms. During the primary bloom, integrated production rates rose to 4 to 4.5 g C m^–2 d^–1, and integrated biomass levels reached 415 to 972 mg chlorophyll m^–2. Primary blooms were usually dominated by large diatoms (Thalassiosira spp.), and in a single year (1989) byChaetoceros spp. The primary blooms terminated upon nutrient depletion in the euphotic zone. Secondary blooms, triggered by nutrient resupply from below, occurred sporadically after the primary bloom and accounted for 4 to 31% of total spring production. The date of initiation and the duration of the primary bloom varied little from year to year (standard deviation 3 and 5 d, respectively). Seasonal production rates and biomass levels varied interannually by a factor of 2 to 3. In contrast, intra-annual variations of more than an order of magnitude, especially in biomass, occurred over periods as short as 10 d. These large variations over short time periods indicate the importance of synchronous timing between spring blooms and the production of larval fish and shellfish, which depend on an appropriate and adequate food supply for growth and survival. Parameters describing primary production (e.g. peak daily production, mean daily production, and total production during the primary bloom and the entire season) exhibited little interannual variation (coefficient of variation, CV = 10 to 19%), but a large degree of intra-annual variation (CV = 77 to 116%). Similarly, interannual variations in biomass (peak chlorophyll, mean chlorophyll) were also lower (CV = 20 to 33%) than intra-annual variations (CV = 85 to 120%).     

The effects of seston food quality on planktonic food web patterns

In planktonic food webs, the conversion rate of plant material to herbivore biomass is determined by a variety of factors such as seston biochemical/elemental composition, phytoplankton cell morphology, and colony architecture. Despite the overwhelming heterogeneity characterizing the plant–animal interface, plankton population models usually misrepresent the food quality constraints imposed on zooplankton growth. In this study, we reformulate the zooplankton grazing term to include seston food quality effects on zooplankton assimilation efficiency and examine its ramifications on system stability. Using different phytoplankton parameterizations with regards to growth strategies, light requirements, sinking rates, and food quality, we examined the dynamics induced in planktonic systems under varying zooplankton mortality/fish predation, light conditions, nutrient availability, and detritus food quality levels. In general, our analysis suggests that high food quality tends to stabilize the planktonic systems, whereas unforced oscillations (limit cycles) emerge with lower seston food quality. For a given phytoplankton specification and resource availability, the amplitude of the plankton oscillations is primarily modulated from zooplankton mortality and secondarily from the nutritional quality of the alternative food source (i.e., detritus). When the phytoplankton community is parameterized as a cyanobacterium-like species, conditions of high nutrient availability combined with high zooplankton mortality led to phytoplankton biomass accumulation, whereas a diatom-like parameterization resulted in relatively low phytoplankton to zooplankton biomass ratios highlighting the notion that high phytoplankton food quality allows the zooplankton community to sustain relatively high biomass and to suppress phytoplankton biomass to low levels. During nutrient and light enrichment conditions, both phytoplankton and detritus food quality determine the extent of the limit cycle region, whereas high algal food quality increases system resilience by shifting the oscillatory region towards lower light attenuation levels. Detritus food quality seems to regulate the amplitude of the dynamic oscillations following enrichment, when algal food quality is low. These results highlight the profitability of the alternative food sources for the grazer as an important predictor for the dynamic behavior of primary producer–grazer interactions in nature.     

The role of ciliates,heterotrophic dinoflagellates and copepods in structuring spring plankton communities at Helgoland Roads,North Sea

Mesocosm experiments coupled with dilution grazing experiments were carried out during the phytoplankton spring bloom 2009. The interactions between phytoplankton, microzooplankton and copepods were investigated using natural plankton communities obtained from Helgoland Roads (54°11.3′N; 7°54.0′E), North Sea. In the absence of mesozooplankton grazers, the microzooplankton rapidly responded to different prey availabilities; this was most pronounced for ciliates such as strombidiids and strobilids. The occurrence of ciliates was strongly dependent on specific prey and abrupt losses in their relative importance with the disappearance of their prey were observed. Thecate and athecate dinoflagellates had a broader food spectrum and slower reaction times compared with ciliates. In general, high microzooplankton potential grazing impacts with an average consumption of 120% of the phytoplankton production (P _p ) were measured. Thus, the decline in phytoplankton biomass could be mainly attributed to an intense grazing by microzooplankton. Copepods were less important phytoplankton grazers consuming on average only 47% of P _p . Microzooplankton in turn contributed a substantial part to the copepods’ diets especially with decreasing quality of phytoplankton food due to nutrient limitation over the course of the bloom. Copepod grazing rates exceeded microzooplankton growth, suggesting their strong top-down control potential on microzooplankton in the field. Selective grazing by microzooplankton was an important factor for stabilising a bloom of less-preferred diatom species in our mesocosms with specific species (Thalassiosira spp., Rhizosolenia spp. and Chaetoceros spp.) dominating the bloom. This study demonstrates the importance of microzooplankton grazers for structuring and controlling phytoplankton spring blooms in temperate waters and the important role of copepods as top-down regulators of microzooplankton.     

Picophytoplankton responses to changing nutrient and light regimes during a bloom

The spring bloom in seasonally stratified seas is often characterized by a rapid increase in photosynthetic biomass. To clarify how the combined effects of nutrient and light availability influence phytoplankton composition in the oligotrophic Gulf of Aqaba, Red Sea, phytoplankton growth and acclimation responses to various nutrient and light regimes were recorded in three independent bioassays and during a naturally-occurring bloom. We show that picoeukaryotes and Synechococcus maintained a “bloomer” growth strategy, which allowed them to grow quickly when nutrient and light limitation were reversed. During the bloom picoeukaryotes and Synechococcus appeared to have higher P requirements relative to N, and were responsible for the majority of photosynthetic biomass accumulation. Following stratification events, populations limited by light showed rapid photoacclimation (based on analysis of cellular fluorescence levels and photosystem II photosynthetic efficiency) and community composition shifts without substantial changes in photosynthetic biomass. The traditional interpretation of “bloom” dynamics (i.e., as an increase in photosynthetic biomass) may therefore be confined to the upper euphotic zone where light is not limiting, while other acclimation processes are more ecologically relevant at depth. Characterizing acclimation processes and growth strategies is important if we are to clarify mechanisms that underlie productivity in oligotrophic regions, which account for approximately half of the global primary production in the ocean. This information is also important for predicting how phytoplankton may respond to global warming-induced oligotrophic ocean expansion.     

The Baltic Sea spring phytoplankton bloom in a changing climate: an experimental approach

The response of the Baltic Sea spring bloom was studied in mesocosm experiments, where temperatures were elevated up to 6°C above the present-day sea surface temperature of the spring bloom season. Four of the seven experiments were carried out at different light levels (32–202?Wh?m^?2 at the start of the experiments) in the different experimental years. In one further experiment, the factors light and temperature were crossed, and in one experiment, the factors density of overwintering zooplankton and temperature were crossed. Overall, there was a slight temporal acceleration of the phytoplankton spring bloom, a decline of peak biomass and a decline of mean cell size with warming. The temperature influence on phytoplankton bloom timing, biomass and size structure was qualitatively highly robust across experiments. The dependence of timing, biomass, and size structure on initial conditions was tested by multiple regression analysis of the y-temperature regressions with the candidate independent variables initial light, initial phytoplankton biomass, initial microzooplankton biomass, and initial mesozooplankton (=copepod) biomass. The bloom timing predicted for mean temperatures (5.28°C) depended on light. The peak biomass showed a strong positive dependence on light and a weaker negative dependence on initial copepod density. Mean phytoplankton cell size predicted for the mean temperature responded positively to light and negatively to copepod density. The anticipated mismatch between phytoplankton supply and food demand by newly hatched copepod nauplii occurred only under the combination of low light and warm temperatures. The analysis presented here confirms earlier conclusions about temperature responses that are based on subsets of our experimental series. However, only the comprehensive analysis across all experiments highlights the importance of the factor light.     

Mass encystment and sinking of dinoflagellates during a spring bloom

The decline of a spring bloom dominated by dinoflagellates and the mass sedimentation of dinoflagellate cysts was documented in a coastal area of the northern Baltic Sea, SW Finland in 1983. The exceptionally large spring phytoplankton bloom was observed in early May. After depletion of nitrate phytoplankton biomass declined rapidly. The bloom was followed by intense sedimentation of spherical cysts and of organic matter at the end of May. These cysts were presumably hypnozygotes of Peridinium hangoei Schiller. Sedimentation of dinoflagellate cysts was estimated to correspond to ca. 45% of the maximum sedimentation of particulate organic carbon at this time, although most of the dinoflagellate biomass disintegrated already in the water column and was deposited as organic detritus or washed away by advection. It is concluded that the life cycle strategies of the dominant vernal phytoplankton species have a major impact on the sedimentation of the spring bloom.     

Succession patterns of phytoplankton blooms: directionality and influence of algal cell size

Using four replicate microcosms in the laboratory, we induced a phytoplankton bloom by enclosing a natural community sampled from Masnou Harbor (N.E. Spain) in November 1987, and examined the pattern of algal succession during the bloom. Good replicability of the temporal patterns of the community biomass and the abundance of most species demonstrated that succession was a directional, non-random process. The successional pathway observed (small flagellates » small centric diatoms » small flagellates) resembled that observed by other authors studying phytoplankton blooms. This pattern differed from previous models of algal succession in that dinoflagellates never comprised a substantial fraction of the community biomass, and in that algal cell size did not tend to increase along the successional sequence. Algal cell size, however, was an important determinant of phytoplankton community structure, since it constrained the density, but not the biomass, achievable by the different species. We suggest that there is not a single, general pattern of phytoplankton succession, but that distinction should be made, at least between seasonal and bloom patterns of phytoplankton succession.     

Modeling the combined effect of nutrients and pyrene on the plankton population: Validation using mesocosm experiment data and scenario analysis

A new integrated model that includes a hydrodynamic model coupled with a contaminant fate and effect sub-model and an ecological sub-model is presented and validated using data from mesocosm experiments. The experiments were carried out in the Isefjord (Denmark) and include the combined effects of nutrients and pyrene addition on the lower trophic levels of bacteria, zooplankton and phytoplankton. The model was able to correctly represent the main dynamics observed in the mesocosms during the 11 days of the experiment and thereby confirmed that it is possible to represent short-term changes in the system with a simplified food-web model on a small spatial and temporal scale. Finally, the validated model was used to carry out a scenario analysis to investigate the effects of a contaminant pulse at different pyrene concentrations and different release timings. Results showed that the ecosystem's vulnerability to a pyrene pulse depends on the initial condition of the system. Stronger biomass reduction was observed when the pulse was released during the zooplankton bloom. Conversely, when the pulse was added at low biomass and before the bloom, the system showed a tendency to behave non-linearly.     

Phytoplankton carbon dynamics during a winter-spring diatom bloom in an enclosed marine ecosystem: primary production,biomass and loss rates

Phytoplankton production, standing crop, and loss processes (respiration, sedimentation, grazing by zooplankton, and excretion) were measured on a daily basis during the growth, dormancy and decline of a winter-spring diatom bloom in a large-scale (13 m³) marine mesocosm in 1987. Carbonspecific rates of production and biomass change were highly correlated whereas production and loss rates were unrelated over the experimental period when the significant changes in algal biomass characteristic of phytoplankton blooms were occurring. The observed decline in diatom growth rates was caused by nutrient limitation. Daily phytoplankton production rates calculated from the phytoplankton continuity equation were in excellent agreement with rates independently determined using standard ¹⁴C techniques. A carbon budget for the winter bloom indicated that 82.4% of the net daytime primary production was accounted for by measured loss processes, 1.3% was present as standing crop at the end of the experiment, and 16.3% was unexplained. Losses via sedimentation (44.8%) and nighttime phytoplankton respiration (24.1%) predominated, while losses due to zooplankton grazing (10.7%) and nighttime phytoplankton excretion (2.8%) were of lesser importance. A model simulating daily phytoplankton biomass was developed to demonstrate the relative importance of the individual loss processes.     

Modelling cyanobacteria in shallow coastal seas

There is an increasing need to describe cyanobacteria bloom dynamics using ecosystem models. We consider two fundamentally different ways how cyanobacteria are currently implemented: a simple approach without explicit consideration of the life cycle which assumes that cyanobacteria grow due to nitrogen fixation alone and an advanced approach that computes the succession of four different stages of the cyanobacteria life cycle based on internal quotas of energy and nitrogen. To qualitatively and quantitatively intercompare these different approaches and with observations, we use the Baltic Sea ecosystem model ERGOM coupled to the one-dimensional water column model GOTM. Four experiments are carried out: three, using the simple approach with either (a) a prescribed constant minimum production, (b) no minimum value or (c) a prescribed constant minimum concentration, and one with (d) the full predictive life cycle. The model data of 35 years (1970-2005) are analyzed for the timing of the bloom, the interannual variability, the annual mean nitrogen fixation rates and the effect of cyanobacteria on eukaryotic phytoplankton. The results show significant differences. In the climatological seasonal mean, only the advanced approach which resolves the life cycle produces a realistic bloom onset and duration. The interannual variability of blooms is unrealistically small in the experiments with a prescribed minimum value. Annual mean nitrogen fixation rates diverge by up to 30% between the four model solutions. Finally, the representation of the cyanobacteria also influences the seasonal cycle of eukaryotic phytoplankton, i.e., flagellates. This study demonstrates that the way how cyanobacteria are implemented in coupled biological-physical models strongly determines the fluxes into the system and between the individual compartments.     

Microbial clumps in seawater in the euphotic zone of Saanich Inlet (British Columbia)

Viable heterotrophic microorganisms were enumerated to be 3.7±7.3 bacterial cells per microbial clump during summer in the euphotic zone of Saanich Inlet, British Columbia, Canada. Large microbial aggregates were observed, especially after the phytoplankton bloom, when the phytoplankton biomass formed about 1/2 the total suspended organic matter in the sea. The cell number per microbial clump was minimal when the phytoplankton fraction in the total suspended organic matter was almost 0 (i.e., before the phytoplankton bloom), and again when the phytoplankton bloom occurred. The size of the microbial clumps is discussed, particularly in reference to the food chain in the sea.The work was carried out at the Fisheries Research Board of Canada, Biological Station, Nanaimo, during the tenure of a National Research Council Post-doctoral fellowship.     

Microphytoplankton in the Strait of Otranto (eastern Mediterranean)

The Strait of Otranto is the connection between the Adriatic and Ionian Seas. Low nutrient concentrations, high transparency, and low phytoplankton cell density and biomass reflect the oligotrophic character of the area. Enrichment of the euphotic layer with nutrients is mainly due to discharge of Albanian and Greek rivers, as well as mixing and upwelling in winter/early spring. Following phytoplankton bloom in April, a progressive decrease of phytoplankton cell density is due to the consumption of nutrients throughout the proceeding summer and autumn. Nitrogen was a strong limiting factor for phytoplankton growth in summer. Deep biomass maxima were detected in the 50 to 100 m layer and corresponded mostly to cells smaller than 20 m. The eastern part of the strait is mostly influenced by the northerly inflowing current from the Ionian Sea, and the western part by the southerly outflowing current from the Adriatic Sea. This typical circulation could be disturbed by inertial oscillations in the current field, generated by the strong oscillating winds and cyclonic eddies. The type of circulation determined the distribution of thermohaline characteristics, abundance, biomass, as well as taxonomic composition of phytoplankton, across the strait. Ecological characteristics of the water masses on two sides of the strait were significantly different during the formation of a longitudinal thermohaline front in May 1990.     

Mechanistic origins of variability in phytoplankton dynamics. Part II: analysis of mesocosm blooms under climate change scenarios

Driving factors of phytoplankton spring blooms have been discussed since long, but rarely analyzed quantitatively. Here, we use a mechanistic size-based ecosystem model to reconstruct observations made during the Kiel mesocosm experiments (2005–2006). The model accurately hindcasts highly variable bloom developments including community shifts in cell size. Under low light, phytoplankton dynamics was mostly controlled by selective mesozooplankton grazing. Selective grazing also explains initial dominance of large diatoms under high light conditions. All blooms were mainly terminated by aggregation and sedimentation. Allometries in nutrient uptake capabilities led to a delayed, post-bloom dominance of small species. In general, biomass and trait dynamics revealed many mutual dependencies, while growth factors decoupled from the respective selective forces. A size shift induced by one factor often changed the growth dependency on other factors. Within climate change scenarios, these indirect effects produced large sensitivities of ecosystem fluxes to the size distribution of winter phytoplankton. These sensitivities exceeded those found for changes in vertical mixing, whereas temperature changes only had minimal impacts.     

Stable isotopes identify an ontogenetic niche expansion in Antarctic krill (Euphausia superba) from the South Shetland Islands, Antarctica

Antarctic krill (Euphausia superba) occupy a key position in the Southern Ocean linking primary production to secondary consumers. While krill is a dominant grazer of phytoplankton, it also consumes heterotrophic prey and the relative importance of these two resources may differ with ontogeny. We used stable isotope analyses to evaluate body size-dependent trophic and habitat shifts in krill during the austral summer around the South Shetland Islands, Antarctica. We found evidence for an asymmetric, ontogenetic niche expansion with adults of both sexes having higher and more variable δ¹⁵N values but consistent δ¹³C values in comparison with juveniles. This result suggests that while phytoplankton likely remains an important life-long resource, krill in our study area expand their dietary niche to include higher trophic food sources as body size increases. The broader dietary niches observed in adults may help buffer them from recent climate-driven shifts in phytoplankton communities that negatively affect larval or juvenile krill that rely predominately on autotrophic resources.     

Physical and biological controls of algal blooms in the Río de la Plata

Coupled three-dimensional hydrodynamic and ecological numerical simulations were used to investigate the role of transport, stagnation zones and dispersion on inter-annual blooms of the diatom Aulacoseira sp. in the vicinity of the drinking water intakes of the Buenos Aires city (Argentina) in the upper Río de la Plata. Three different summer events were analyzed. First, a mild biomass bloom year (2006–2007), second, a high biomass bloom year (2007–2008) and third, a “normal” no bloom year (2009–2010). Simulated water height, water temperature, suspended solids and chlorophyll \(a\) concentrations patterns compared well with field data. Results revealed that the advection of phytoplankton cells via inflows to the Río de la Plata triggered Aulacoseira sp. blooms in the domain. In addition, excessive growth observed near the drinking water intakes, along the Argentinean margin, were associated with long retention times (stagnant region) and weak horizontal dispersion. Increased concentrations of suspended solids in the water column, in response to re-suspension events, did not prevent the blooms, however, were found to also play a key role in controlling the rate of phytoplankton growth. Finally, a non-dimensional parameter, R, that considers phytoplankton patch size, e-folding growth and dispersion time scales is shown to determine the potential bloom occurrences, as well as bloom intensity; R values higher than 5.7 suggest intense phytoplankton growth. For the mild biomass bloom year, \(R = 7.5\) , for the high biomass bloom year, \(R = 11\) and for the “normal” no bloom year \(R= 0.4\) .     

Seasonal studies on the relative importance of different size fractions of phytoplankton in Narragansett Bay (USA)

The composition and productivity of four different size-fractions (<20, 20 to 60, 60 to 100, >100 μm) of the phytoplankton of lower Narragansett Bay (USA) were followed over an annual cycle from November, 1972 to October, 1973. Diatoms dominated the population in the winter-spring bloom and in the fall, the summer population was dominated by flagellates. The nannoplankton (<20 μm) were the most important, accounting for 46.6% of the annual biomass as chlorophyll a and 50.8% of the total production. The relative importance of the different fractions showed a marked seasonality. During the winter-spring and fall blooms the netplankton fractions (>20 μm) were the most important. Nannoplankters domnated in the summer. The yearly mean assimilation numbers for the different fractions were not signfficantly different. During the winter-spring bloom, however, the assimilation numbers for the netplankters were significantly higher than those for the nannoplankton fraction. Temperature accounted for most of the variability in assimilation numbers; a marked nutrient stress was observed on only two occasions. Growth rates calculated from ¹⁴C uptake and adenosine triphosphate (ATP)-cell carbon were generally quite high; maxima were >1.90 doublings per day during blooms of a flagellate in the summer and of Skeletonema costatum in the fall. The series of short cycles observed in which the dominant species changed were related to changes in the physiological state of the population. Higher growth rates were generally observed at times of peak phytoplankton abundance while lower growth rates were observed between these peaks. The high growth rates and assimilation numbers usually found suggest that the phytoplankton in lower Narragansett Bay was not generally nutrient-limited between November, 1972 and October, 1973. Nutrient regeneration in this shallow estuary, therefore, must be very rapid when in situ nutrient levels are low.     

Distribution and structure of heterotrophic protist communities in the northeast equatorial Pacific Ocean

The distribution and structure of heterotrophic protist communities and size-fractionated chlorophyll a were studied during the Korea Deep Ocean Study 98 (KODOS 98) research expedition (July 1998) in the northeast equatorial Pacific Ocean (5–11°N). Areas of convergence and divergence formed at the boundaries of the South Equatorial Current (SEC), North Equatorial Current (NEC), and North Equatorial Counter Current (NECC) during the expedition. Water column physicochemical characteristics significantly influenced the size structure of heterotrophic protist communities. Intense vertical mixing and high nutrient and chlorophyll a concentrations characterized SEC and NECC areas, which were affected by converging and diverging water masses, respectively. Nanophytoplankton dominated in SEC and NECC areas; both areas also had relatively high heterotrophic protist biomasses (average 743 µg C m^–2). NEC areas were characterized by a stratified vertical structure, low nutrient and chlorophyll a concentrations, and picophytoplankton dominance. The heterotrophic protist biomass in NEC areas averaged 414 µg C m^–2; nanoprotists (<20 µm) dominated the community. The nanoprotist biomass comprised 49–54% of the total heterotrophic protist biomass in SEC/NECC areas and 67–72% in NEC areas. The biomass of heterotrophic protists was higher in SEC/NECC areas than in NEC areas, but the relative importance of nanoprotists was greater in NEC areas than in SEC/NECC areas. Heterotrophic dinoflagellates were dominant components of the <20 µm and >20 µm size classes in both water columns. The biomass of heterotrophic protists significantly correlated with the net-, nano-, and picophytoplankton biomass in SEC/NECC areas and with the nano- and picophytoplankton biomass in NEC areas. Heterotrophic protists and phytoplankton also showed strong positive correlation in the study area. The size structure of the phytoplankton biomass coincided with that of heterotrophic protists; the heterotrophic protist biomass positively correlated with the protists prey source. These relationships suggest that the community structure of heterotrophic protists and the microbial food web depended on size classes within the phytoplankton biomass. Microzooplankton grazing and phytoplankton growth rates were higher in SEC/NECC areas than in NEC areas. In contrast, the potential primary production grazed by microzooplankton was relatively high in NEC areas (127.3%) compared with SEC/NECC areas (94.6%). Our results indicate that the relative importance and size structure of heterotrophic protists might vary according to two distinct water column structures.Communicated by T. Ikeda, Hakodate     

Simple predator–prey interactions control dynamics in a plankton food web model

A plankton food web model is analysed using interaction parameter values appropriate to the upper mixed layer of the high latitude oceans. The dynamics of this four-variable system are analysed in terms of the dynamics of much simpler two-variable predator–prey subsystems. Thus, the food web's robust, periodic, four-dimensional dynamics are explained by means of two-dimensional spirals and limit cycles. These dynamical subsystems are coupled by means of an omnivore that transfers control of the dynamics between the two predator–prey subsystems. The food web may substantially decouple the predator–prey subsystems so that the oscillating phytoplankton/zooplankton blooms exhibit population collapses when bacterial ‘breathers’ briefly dominate after growing dramatically from low background levels. This regular bloom/breather behaviour becomes benignly chaotic when the system is mildly forced by the annual cycle of the sun's irradiance.     

Grazing regulates the spatial variability of periphyton biomass

The presence of consumers not only alters the mean biomass of the prey assemblage, but also affects the spatial heterogeneity of biomass distribution. Whereas the mean prey biomass is generally reduced by consumer presence, the effect on spatial heterogeneity is less clear-cut. A meta-analysis of almost 600 field experiments manipulating the presence of benthic invertebrate or vertebrate grazers was conducted to analyze the effect of grazers on both the absolute spatial variability of periphyton biomass and the relative variability, which was standardized to the mean. Effects on absolute variability were measured as the log response ratio of the standard deviation of biomass (LR-SD), whereas effects on relative variability were measured as the log response ratio of the coefficient of variation of biomass (LR-CV). The overall magnitude and range of LR-SD and LR-CV indicated that grazers not only reduced periphyton biomass, but also substantially altered their spatial distribution. However, grazer effects differed strongly for absolute and relative variability. On average, grazers reduced the absolute spatial variability in prey biomass by 50% (average LR-SD = -0.68) but increased the relative variability by 24% (average LR-CV = 0.22). The magnitude of LR-SD strongly depended on the efficiency of grazing, with strong biomass removal leading to strong homogenization. Moreover, LR-CV and LR-SD were significantly affected by habitat type (freshwater vs. coastal) and substrata. Given the importance of spatial heterogeneity for resource uptake, competition and the maintenance of diversity, grazer presence has potentially strong indirect effects on the interactions within prey assemblages.     

Sexuality and cyst formation of the spring-bloom dinoflagellate Scrippsiella hangoei in the coastal northern Baltic Sea

The temporal sequence and the magnitude of the sexual reproduction and subsequent cyst deposition of the common spring-bloom dinoflagellate Scrippsiella hangoei (Schiller) Larsen was studied during spring 1996 on the SW coast of Finland, Baltic Sea. The abundances of the different size of fractions of S. hangoei (14 to 18 μm, 18 to 22 μm and >22 μm) were monitored in the water column, and the deposition of resting cysts was measured using moored sediment traps. Cyst sedimentation rates were measured throughout the seasonal cycle in order to estimate cyst resuspension rates for the quantitative assessment of the fraction of population undergoing encystment. The onset of sexual reproduction, indicated by a significant increase of the small cells (14 to 18 μm) representing gametes, occurred in a nutrient replete environment well before the exponential growth phase and peak abundances of vegetative cells. Gamete formation was followed by high abundances of large cells (>22 μm) representing planozygotes, and subsequent sedimentation of resting cysts. Approximately 60% of the asexually growing bloom population was estimated to form planozygotes, suggesting that encystment was an important factor in bloom termination and possibly plays a role in the regulation of the magnitude of the bloom. Finally encystment accounted for 40% of the entire S. hangoei population, resulting in a considerable loss of the bloom population and an input of the vernal phytoplankton biomass to the benthos. Received: 11 December 1998 / Accepted: 8 April 1999