The phased division of the freshwater dinoflagellate Ceratium hirundinella and its use as a method of assessing growth in natural populations

SUMMARY. The nocturnal phasing and partial synchrony of cell division in Ceratium hirundinella was investigated on four occasions for dense planktonic populations in a small productive lake (Esthwaite Water). The population growth rates deduced from the proportion of cells dividing per day are compared with the rates of increase of cell density in the lake. The maximum proportion of Ceratium cells found dividing at any time was 5.8 ± 1.0%, and the time of optimum division was 03.00 hours G.M.T. The daily rate of division during the main phase of population increases was similar to that deduced from the overall population increments at that time, although during the week of collection the increase had apparently ceased temporarily. On the other three occasions, either increase of cell numbers had ceased or the population was declining, but a continued low rate of division ( c . 3% day⁻¹) was found.
The nocturnal division of Ceratium in Esthwaite Water is compared with the division phasing of dinaflagellates described from elsewhere. Some general problems associated with the derivation of estimates of population growth rate from division frequency are also considered.     

Vertical microscale patchiness in nano- and microplankton distributions in a stratified estuary

Microscale (decimetre) vertical heterogeneity in the distributionof nano- and microplankton was studied on August 23, 1999 attwo different sites (separated by ~40 m) in Aarhus Bay, Denmark.At the time of sampling, the water column was stratified withrespect to both temperature and salinity and a subsurface fluorescencemaximum (corresponding to ~10 µg chlorophyll a l^-1) occurredimmediately below the primary pycnocline (9 m). Samples forspecies identification were taken at the surface, near the bottomof the water column and at 20 15-cm intervals in and aroundthe depth of maximum fluorescence. The plankton communitiesrecorded in the three different regions of the water columndiffered dramatically from one another. In addition, significantdifferences were found in the distribution patterns of speciesand functional groups in the region of sampling around the fluorescencepeak. The same patterns in vertical species distributions wereobserved at the two stations. In the region surrounding thesubsurface fluorescence peak, diatoms were, generally, regularlydistributed, although the total diatom biomass decreased slightlywith depth. Dinoflagellate species were mainly non-regularlydistributed and could be divided into two groups: (i) autotrophicor potentially mixotrophic species (Dinophysis norvegica, Dinophysisacuminata, Dinophysis cf. dens, Prorocentrum micans, Gymnodiniumchlorophorum and Ceratium macroceros) that mainly decreasedin numbers with depth or were aggregated in their distribution;and (ii) heterotrophic or potentially mixotrophic species (Ceratiumlineatum, Ceratium longipes, Ceratium furca, Ceratium tripos,Ceratium fusus, Protoperidinium curtipes, Protoperidinium steinii,Diplopsalis spp. and Katodinium glaucum). In the latter group,the species mainly increased in numbers with depth or were randomlydistributed. Most ciliates were uniformly distributed verticallyin the water column. However, small cells of the photosynthesizingciliate Mesodinium rubrum were most abundant at the depth ofmaximal fluorescence while large M. rubrum cells were equallyabundant at all the depths sampled, suggesting the two sizegroups of this organism may differ ecologically. Overall, thestudy demonstrates that traditional plankton sampling methodsmay lead to misinterpretations of speciesco-existence and interactions.     

The horizontal distribution of plankton in a deep, oligotrophic lake—Loch Ness, Scotland

1. The horizontal distribution of plankton in Loch Ness, a deep, oligotrophic lake with a simple trench morphometry, was studied on three occasions in 1993. Samples were collected from ten stations spaced along the length of the loch and the abundance of algae, bacteria, protozoa, rotifers and crustacean zooplankton was determined. 2. Horizontal variability was found to be greatest for the metazoan zooplankton and for the algae, especially two cryptomonad flagellates. Bacteria and heterotrophic protozoa showed relatively little horizontal variability. The degree of horizontal variability was not sufficient seriously to affect studies of the seasonality of plankton abundance conducted from a single sampling station. 3. Gradients of plankton distribution along the length of the loch were dependent on the recent wind history. The direction of the gradient could readily reverse with a shift in wind direction. These results indicate that the horizontal distribution of plankton in Loch Ness is more dependent on wind-induced water circulation patterns than on differential growth of plankton in water masses of differing chemistry.     

The role of macroinvertebrates and fish in regulating the provision by macrophytes of refugia for zooplankton in a warm temperate shallow lake

1. The zooplankton often undergoes diel horizontal migration (DHM) from the open water to the littoral of shallow lakes, thus avoiding predators in the former. This behaviour has functional impacts within the lake, as it enhances zooplankton survival, increases their control of phytoplankton and tends to stabilise the clear water state. However, most of the evidence supporting this migration pattern comes from cold north temperate lakes, and more evidence from tropical and subtropical areas, as well as from southern temperate areas, is needed. 2. We conducted a field study of the diel horizontal and vertical migration of zooplankton, and the horizontal distribution of potential predatory macroinvertebrates and fish, over two consecutive days in the summer in a temperate lake in the southern hemisphere. We took zooplankton samples at two depths, at three sampling stations (inside beds of aquatic macrophytes, at their edge and in open water) along three transects running from the centre of a bed of Ceratophyllum demersum to open water. At each sampling station, we also took samples of macroinvertebrates and fish and measured physical and chemical environmental variables. 3. Zooplankton (pelagic cladocerans, calanoid copepods and rotifers) avoided the shore, probably because of the greater risk from predators there. Larger and more vulnerable cladocerans, such as Diaphanosoma brachyurum and Moina micrura, were two to four times more abundant in open water than at the edge of or inside beds of macrophytes, respectively, by both day and night. Less vulnerable zooplankton [i.e. of medium body size (Ceriodaphnia dubia) or with the ability to swim fast (calanoid copepods)] were distributed evenly between open water and the edge of the plant beds. Small zooplankton, Bosmina huaronensis and pelagic rotifers, showed an even distribution among the three sampling stations. Accordingly, no DHM of zooplankton occurred, although larger organisms migrated vertically inside C. demersum stands. 4. Macrophytes contained high densities of predatory macroinvertebrates and fish. The predator assemblage, composed of large‐bodied macroinvertebrates (including odonates and shrimps) and small littoral fish, was permanently associated with submerged macrophytes. None of these groups moved outside the plant beds or changed their population structure (fish) over the diel cycle. 5. Submerged macrophyte beds do not represent a refuge for zooplankton in lakes where predators are numerous among the plants, implying a weaker top‐down control of phytoplankton biomass by zooplankton and, consequently, a more turbid lake. The effectiveness of macrophytes as a refuge for zooplankton depends on the associated assemblage of predatory macroinvertebrates and fish among the plants.     

Spatial patterns of planktonic microcrustaceans in a small shallow lake

The horizontal distribution of planktonic microcrustaceans was studied at two different spatial scales in a small shallow lake (Créteil Lake, Paris suburb, France) during the summer.At all scales of sampling, the distribution was patchy. However, the variations between sampling units were smallest when considering the smallest sampling grid (30 sampling units in a 0.5 ha area). Positive correlations between densities of most taxa were observed at this scale, indicating that a common set of factors could influence zooplankton patchiness.Physical factors, such as wind-induced currents, strongly influenced the lakewide distribution of zooplankton (43 sampling units in a 42 ha area). The horizontal pattern of each taxon appeared to be linked to its ability to form vertical aggregations. Moreover, offshore-nearshore differences in fecundity of several taxa suggest the action of biotic factors in the horizontal distribution of zooplankton. A higher predation risk in the littoral neighbourhood seems to be the more probable proximate - or ultimate - factor which could explain this pattern.In this small and shallow lake, most factors that can affect distribution of zooplankton are temporally variable and unpredictable. In the absence of automatic and efficient measuring techniques, the realisation of a small number of integrated samples at the level of the whole ecosystem could be a simple and suitable solution to obtain statistically reliable data on zooplankton demography.     

A laboratory simulation of diel changes affecting cellular composition of the dinoflagellate Ceratium hirundinella

SUMMARY. Changes of cellular composition of a planktonic dinoflageilate, Ceratium hirundinella , were studied over 2–1–48-h periods in the laboratory on three separate occasions. Samples from a lake population in the stationary phase of the population cycle were incubated at lake temperature under continuous illumination and in a light-dark (LD) regime of LD 14:10.
The magnitudes of diel variability in ratios of carbohydrate:chlorophyll- a , carbohydrate:carbon. carbon:nitrogen and carotenoids:chlorophyll- a for samples incubated at LD 14:10, were similar to those observed in situ at a similar phase of the population growth cycle. The ratios involving carbohydrate showed the most diel variation, which was not reduced under continuous illumination.
By appearance 'dark' and 'pale' cells of Ceratium were distinguished in the fixed samples; a persistent periodicity for the interconversion of dark and pale cells is described. The periodicity and amplitude of inter-conversions were not always in phase with those of chlorophyll- a synthesis and decomposition. The observed changes were almost certainly independent of the light-dark cycle: they were apparently induced by nutrient deficiency.     

Spatial variability in the abundance and composition of the free-living bacterioplankton community in the pelagic zone of Lake Bourget (France)

Spatial variations in the abundance and diversity of the free-living bacterioplankton community of a large Alpine lake, Lake Bourget (France), were investigated in the pelagic zone by means of two two-dimensional samplings taken in 2003. Lake-water samples were collected in winter during water mixing, and in early summer during stratification. The population abundance in each sample was determined by flow cytometry. Denaturing gradient gel electrophoresis of PCR-amplified 16S rRNA gene fragments from organisms measuring less than 2 mum was used to assess eubacterioplankton community composition. In winter, no obvious differences were observed in either the abundance or the diversity of the bacterial community, on either the horizontal or the vertical scales. The only influence detected was that of river water input, but this was at a very minor scale relative to the surface area of the lake. In early summer, changes were found in the community composition on the vertical scale related to the thermal stratification of the water column. There were also marked differences on the horizontal scale at 15 m depth due to internal waves. The implications of these findings for sampling strategies are very important from the perspective of comparative studies of free-living bacterial community diversity and functioning in large and deep lakes.     

Use of time-series analysis to demonstrate advection rates of different variables in a small lake

Time-series analysis of daily data from 3 stations (maximumseparation – 1 km) in a small lake (Guelph Lake, Ontario)has demonstrated the existence of (0 strong horizontal advectionwithin the basin at a time scale of 2–3 d and (ii) longerterm periodicities (10–20 d) associated with the passageof atmospheric weather systems. Different rates and directionsof horizontal advection were observed for different variables.Fluctuations in integral SRP concentrations at station 1 (outflowend) consistently occurred 3 d later than at station 3 (inflowend). The algal biomass (chlorophyll a) at station 1 laggedthat at station 3 by half a day. The time lag between station1 and station 3 for Aphanizomenon flos-aquae was 7 d, whereasfor Ceratitun hirundinella station 1 led station 3 by 3 d indicatingmotion by the species in opposite directions. The distributionof the algal biomass in space and time was a composite of theindividual species patterns. Longer term (10–20 d) periodicitieswere also observed. Examination of the periodicities in theaverage daily wind speed and water column stability showed thatsuch lags or leads were likely due to periods of vertical mixing.The significance of these results is then discussed in relationto sampling schemes for the analysis of phytoplankton dynamicsin small basins.     

Physiological and environmental constraints in the ecology of the planktonic dinoflagellate Ceratium hirundinella

SUMMARY. Constraints upon the vertical distribution of a population of Ceratium hirundinella , in a productive English lake during the summer stratification of 1976, are considered. They are interpreted in relation to vertical gradients of temperature, dissolved oxygen and irradiance, and to rates of photosynthesis and respiration measured as oxygen exchange in long- and short-term exposures. The motile cells tended to aggregate at an intermediate depth in the epilimnion, associated with a relative irradiance level of c . 10% or c . 140 μ einsteins m⁻²s⁻¹ as measured with a horizontal PAR sensor. Higher irradiances, and conditions below the oxycline, were apparently unfavourable, but the intervening layer was severely compressed at the height of summer stratification, when concentrations of inorganic nitrogen were also minimal. Thus the population apparently passed through a critical period, at which the cellular content of chlorophyll-α was much reduced. The preferred irradiance level in the lake corresponded to that at which measured rates of net photosynthesis were maximal. Increased rates of oxygen evolution were measured at higher irradiances in very short exposures; this behaviour may be of importance to cells experiencing vertical movement in nature.     

The spatial distribution of nutrients in the South Basin of Windermere

SUMMARY. The vertical and horizontal distribution of electrical conductivity, soluble reactive phosphorus (SRP), total phosphorus, ammonia-nitrogen and nitrate-nitrogen in the South Basin of Windermere is described and related, where possible, to spatial variations in phytoplankton and zooplankton. For most variables, the maximum range of horizontal variation on a single day was greater than the maximum range of seasonal variation measured at a single station over a full year. Horizontal variations in SRP and ammonia were particularly high, with coefficients of variation often exceeding 100%. The errors associated with sampling at a single station were most pronounced when local accumulations of zooplankton or phytoplankton occurred in the lake. Horizontal variations in conductivity were primarily influenced by the discharge of treated sewage effluent into the central region and the mass transport of water from the more oligotrophic North Basin. Occasionally, more widespread variations in conductivity appeared to be related to spatial variations in photosynthetic activity. The major factor influencing the horizontal distribution of SRP was the discharge of treated effluent into the central region. Despite the intensive recycling of phosphorus, SRP concentrations were nearly always 10–30% higher near the sewage works than elsewhere in the basin. On a few occasions, significant horizontal differences in SRP concentration were also detected within downwind accumulations of crustacean zooplankton. Horizontal variations in total phosphorus were produced by spatial differences in SRP concentration or by local accumulations of phytoplankton or zooplankton. Horizontal differences in ammonia concentration appeared to be produced, both by the effluent discharge and by the turbulent transfer of nutrients from sediments in shallow water. Observations of effluent movement and dispersion demonstrate that wind-induced water movements tend to recirculate nutrient-rich water in the central region and limit mixing along the axis of the basin. The statistical implications of nutrient heterogeneity are discussed in relation to sampling strategy and the possible effects of persistent nutrient concentration gradients on phytoplankton patch formation are assessed.     

Quantification of biogenic 3-D structures in marine sediments

Burrow and tube structures of marine infauna were quantified in sediments from cores obtained at 3 depths (75, 95 and 118 m) at 3 occasions over a 1-year period. The benthic communities at the two deeper stations were re-establishing subsequent to re-oxygenation after hypoxic conditions; the shallow station was a reference station unaffected by hypoxia. The benthic macrofauna was simultaneously quantified from sieved grab samples. The sediment cores were frozen and later sliced with a plane in horizontal layers. Biogenic structures in these sections were digitally analysed for numbers, area and volume. Number of tubes and burrows showed a general decline with depth in the sediment. The biogenic structures in the sediment at the reference station increased the sediment-water interface about 1.5 times compared to the surface area. The surface areas of burrows and tubes were 2 to 3 times larger at the reference station than at the deeper stations. The benthic fauna at the 95-m and 118-m stations showed a temporal increase in density and number of species. In contrast, development of biogenic surface areas in the sediment at these stations showed no significant temporal trend during the benthic faunal succession. Relic tubes of Melinna cristata were found throughout the sampling period at the 118-m station. The ecological importance of bioirrigation for biogeochemical processes is discussed.     

Modelling spatial distributions of Ceratium hirundnella and Microcystis. in a small productive British lake

The short-term relationships between the spatial distributions of phytoplankton and the environmental conditions of Esthwaite Water, a small eutrophic lake in the English Lake District, UK, were examined using a hydrodynamic model. Spatial distributions of phytoplankton were simulated on two occasions the first, when the population was dominated by dinoflagellates; and the second, when the population was dominated by cyanobacteria.Vertical motility of the dinoflagellate Ceratium hirundinellaand buoyancy of the cyanobacteria Microcystis ssprm.were estimated as functions of irradiance. Water velocity fields were estimated through solving the 3-D Navier–Stokes equations on a finite-volume, unstructured non-orthogonal grid. Simulated circulation patterns of water and phytoplankton were similar to those obtained through field observations. Near-surface drift currents were initiated by wind stress, which then generated return currents along the seasonal thermocline. Aggregations of motile Ceratiumthat existed near the thermocline were pushed upwind by the deep return currents and accumulated at upwelling areas. In contrast, near-surface aggregations of Microcystiswere pushed downwind by the surface currents and accumulated at downwelling areas. Horizontal and vertical phytoplankton distributions resulted from the interaction between the vertical motility of the phytoplankton (dependent upon the light environment) and the velocity vectors at the depths at which the phytoplankton accumulated (dependent upon wind stress and morphometry). Modelling showed that phytoplankton motility and buoyancy greatly affect phytoplankton spatial distributions.     

Distribution patterns of zooplankton in Tjeukemeer, The Netherlands

The vertical and horizontal distribution of Copepoda and Cladocerain Tjeukemeer were studied. Indices of dispersion are discussed,the sample variance of log-transformed data proved to be anappropriate way to quantify patchiness. Indices of dispersionwere calculated to express species specific differences. Onlyduring short periods of the year the population densities appearedto be influenced by canals and pumping stations that bring waterto the lake. Dispersion values were influenced by windspeed,inhomogenities in the chlorophyll-a concentrations and by thepopulation densities of the populations studied. Sample variancesare relatively low, compared with those of other lakes.     

基于水声学方法的天目湖鱼类季节和昼夜空间分布研究

采用水声学方法系统地对天目湖春夏秋冬四季鱼类水平和垂直空间分布进行了调查研究,同时对昼夜探测的差异性进行了比较,并对不同季节间鱼类聚群形态进行了探讨。研究结果表明,天目湖鱼类在不同季节间存在规律性水平迁移,从全湖角度分析,天目湖鱼类资源昼夜间的水平空间分布无明显差异特征,但在春末和夏季的局部区域里,昼夜间鱼类存在一定近岸—远岸的水平迁移;不同季节和昼夜间,鱼类垂直分布差异明显,且在存在温跃层的夏季7月昼间,鱼类密度垂直分布与水温的垂直分布关系密切,温跃层以下的鱼类密度基本为0;天目湖鱼类在四个季节都属于成群分布类型,但季节间鱼类聚群形态不同,在冬季的1月呈现出典型的聚集,相对于昼间,春夏季鱼类在夜间分布更为均匀、分散,且水平探测表明在夜间水体表层存在大量鱼类分布。     

Impact of internal waves on the spatial distribution of Planktothrix rubescens (cyanobacteria) in an alpine lake

The vertical and horizontal distribution of the cyanobacterium, Planktothrix rubescens, was studied in a deep alpine lake (Lac du Bourget) in a 2-year monitoring program with 11 sampling points, and a 24-h survey at one sampling station. This species is known to proliferate in the metalimnic layer of numerous deep mesotrophic lakes in temperate areas, and also to produce hepatotoxins. When looking at the distribution of P. rubescens at the scale of the entire lake, we found large variations (up to 10 m) in the depth of the biomass peak in the water column. These variations were closely correlated to isotherm displacements. We also found significant variations in the distribution of the cyanobacterial biomass in the northern and southern parts of the lake. We used a physical modeling approach to demonstrate that two internal wave modes can explain these variations. Internal waves are generated by wind events, but can still be detected several days after the end of these events. Finally, our 24-h survey at one sampling point demonstrated that the V1H1 sinusoidal motion could evolve into nonlinear fronts. All these findings show that internal waves have a major impact on the distribution of P. rubescens proliferating in the metalimnic layer of a deep lake, and that this process could influence the growth of this species by a direct impact on light availability.     

Habitat selection and diel distribution of the crustacean zooplankton from a shallow Mediterranean lake during the turbid and clear water phases

1. The fish fauna of many shallow Mediterranean Lakes is dominated by small‐bodied exotic omnivores, with potential implications for fish–zooplankton interactions still largely unknown. Here we studied diel variation in the vertical and horizontal distribution of the crustacean plankton in Lake Vela, a shallow polymictic and eutrophic lake. Diel sampling was carried out on three consecutive days along a horizontal transect, including an open‐water station and a macrophyte (Nymphaea alba) bed. Since transparency is a key determinant of the predation risk posed by fish, the zooplankton sampling campaigns were conducted in both the turbid (autumn) and clear water (spring) phases. 2. In the turbid phase, most taxa were homogeneously distributed along the vertical and horizontal axes in the three consecutive days. The only exception was for copepod nauplii, which showed vertical heterogeneity, possibly as a response to invertebrate predators. 3. In the clear water phase, most zooplankton taxa displayed habitat selection. Vertically, the general response consisted of a daily vertical migration (DVM), despite the limited depth (1.6 m). Horizontally, zooplankters showed an overall preference for the pelagic zone, independent of the time of the day. Such evidence is contrary to the postulated role of macrophytes as an anti‐predator refuge for the zooplankton. 4. These vertical (DVM) and horizontal (macrophyte‐avoidance) patterns were particularly conspicuous for large Daphnia, suggesting that predation risk from size‐selective predators (fish) was the main factor behind the spatial heterogeneity of zooplankton in the spring. Thus, the difference in the zooplankton spatial distribution pattern and habitat selection among seasons (turbid and clear water phases) seems to be mediated the predation risk from fish, which is directly related to water transparency. 5. The zooplankton in Lake Vela have anti‐predator behaviour that minimises predation from fish. We hypothesise that, due to the distinct fish community of shallow Mediterranean lakes, aquatic macrophytes may not provide adequate refuge to zooplankters, as seen in northern temperate lakes.     

A fish-kill in Heart Lake, Ontario, associated with the collapse of a massive population of Ceratium hirundinella (Dinophyceae)

SUMMARY. A massive population of the common dinoflagellate Ceratium hirundinella developed in Heart Lake. Ontario, Canada during the summer of 1976 and its sudden collapse and subsequent decomposition depleted dissolved oxygen and resulted in a fish-kill in the lake. The lake was being artificially mixed at the time by supplying compressed air to the bottom waters and the limnological events contributing to the development of the Ceratium population and its collapse appear to be closely related to the artificial destratification process. Artificial destratification during 1976 precluded the development of blue-green algue. The process also led to an increase in the density of herbivorous zooplankters which controlled the development of smaller planktonic algae. Ceratium flourished in Heart Lake because there was little competition for nutrients from other algae and because Ceratium cells are too large to be grazed by the zooplankton. The maximum size of the Ceratium population (53 mm³ 1⁻¹) is apparently the highest biomass reported in the literature and its collapse may have been related to a depletion of inorganic nitrogen. There is apparently no previously published record of a Ceratium -induced fish-kill in a freshwater lake.     

Temporal and spatial variation of coastal zooplankton in the Baltic Sea

When analysing temporal variation monthly mean abundances of zooplankton sampled at a eoastal station in the northern Baltic Sea between 1976 and 1988 showed the lowest between-year variation in the summers The coefficients of variation were estimated at 50% for rotifers in June. 70% for cladocerans in August and between 30 and 50% for different copepodite stages in August Moreover in the summers, all dominating zooplankton groups were abundant Estimates of yearly biomass or production of zooplankton must, however, be based on frequent sampling during the whole year because of large differences between months in zooplankton composition and abundance
In 1989. spatial variation was analyzed by sampling zooplankton on three occasions along a 9 8 km transect with 15 stations, 700 m apart All taxonomic/life-stage groupings of zooplankton were heterogeneously distributed Geostatistical analysis did not give a definite answer to what the minimum distance between stations should be to obtain independent samples but indicated that 700 to 1400 m would be sufficient in most cases For most of the zooplankton groupings the coefficients of variation in the sampled area were estimated at 20-40% on all sampling occasions     

千岛湖浮游动物的群落结构

研究了贫－中营养型的大型,深水湖泊－－浙江千岛湖浮游动物的群落结构,包括种类组成,种群动态,现存量及群落多样性指数。在1a的研究中,共发现139种浮游动物（27种原生动物,70种轮虫,26种枝角类和16种桡足类）。根据年平均密度,各类浮游动物的优势种分别为褶累枝(Epistylis plicatilie),螺形龟甲轮虫9Keratella cochlearis),透明蚤（Daphnia hyalina)和一种中剑水蚤（Mesocyclops notius)。轮虫和枝角类群落多样性指数随着水体透明度的增大呈下降的趋势;采样站变异下,枝角类群落多样性指数与其种类数和密度呈显著的正相关关系;月份变异下,桡足类群落多样性指数与其密度呈显著的正相关关系。     

Spatial distribution of motile phytoplankton in a stratified reservoir: the physical controls on patch formation

Changes in the spatial distribution of the dinoflagellate Ceratiumhirundinella were observed in a stratified, medium-sized (16km²) Argentinean reservoir over several days. A fluorescenceprofiling technique was used to identify persistent patchinessin the distribution of the dinoflagellate. A three-dimensionalnumerical model was used to reconcile a range of different unsteadyprocesses and prove that the initial source of heterogeneityin the system was the vertical migration of Ceratium. Once migrationestablished vertical heterogeneity, the dominant influence onthe patch dynamics alternated between control by migration andcontrol by mixing and transport. This led to the developmentof persistent horizontal patchiness. The analysis revealed thatthe region of the lake inhabited by Ceratium was highly predictableand from this result it was determined that physical processes(with some influence from migration) control the habitat ofthis dinoflagellate rather than biological/chemical gradients.When the spatial habitat of a particular phytoplankton speciescan be isolated in this manner, the resources available to thespecies can be more accurately determined by further study.The results are particularly applicable to the study of motile/buoyantplankton in aquatic systems that are periodically subject tomoderate or strong wind forcing events.