Protozooplankton in acidic mining lakes with special respect to ciliates

The planktonic protozoa, especially ciliates, were analysed in five mining lakes of various pH and acidity values in the Lusatian and mid-German mining area. Heliozoa were the main protozoan component in the very low pH (<2.9) lakes. In the lakes with pH >2.9, the ciliate community consisted of Hypotrichida, Prostomatida and Peritrichida. The species diversity of the ciliate community was reduced and typical representatives of plankton ciliates were lacking. During periods when populations were at their peak, the cell numbers and biomass of ciliates were comparable to those neutral lakes, reaching maximum values of 30 000 cells l^–1 and 0.3 mg l^–1. Cell numbers and biomass peaks appeared without a clear seasonal pattern but in some cases a correlation to phytoplankton dynamics was recognizable.     

Vertical and Seasonal Dynamics of Planktonic Ciliates in a Strongly Stratified Hypertrophic Lake

Seasonal population dynamics and the vertical distribution of planktonic ciliates in a hypertrophic and strongly stratified temperate lake were studied from April to October in 2000 and from April to June in 2001. In the epi- and metalimnion the ciliate abundance peaked in spring and late summer, reaching maximum values in the metalimnion (86 cells ml⁻¹) on 7th August 2000. In the epilimnion, the highest biomass content (414 μg C l⁻¹) was observed on 8th May 2000. In the hypolimnion only a late summer peak occurred and the ciliate numbers were always lower than in the epi- and metalimnion. Five groups dominated the community of ciliates: Oligotrichida, Gymnostomatea, Prostomatida, Hymenostomata and Peritrichia, and the community composition varied greatly with depth. In the epilimnion the ciliate numbers were dominated by oligotrichs but small algivorous prostomatids, peritrichs and gymnostomes were also numerous. In the metalimnion these groups were gradually replaced by scuticociliates and mixotrophic Coleps spp. In the hypolimnion scuticociliates and species known as benthic migrants dominated. In the epilimnion and upper metalimnion in spring large herbivores and in summer small bacterivores were more numerous.     

Vertical distribution of planktonic ciliates in strongly stratified temperate lakes

The vertical distribution of planktonic ciliates in eight strongly stratified temperate lakes was studied in summer 1998. Ciliate abundance and biomass were highest (mean 39.9 cells ml^–1 and 181.9 g C l^–1) in the epi-, and lowest (mean 8.2 cells ml^–1 and 97.6 g C l^–1) in the hypolimnion. The community of ciliates was dominated by five orders: Oligotrichida, Haptorida, Prostomatida, Scuticociliatida and Peritrichida. The community composition varied greatly with depth. In the epilimnion, the ciliate numbers were dominated by oligotrichs but small algivorous prostomatids, peritrichs and haptorids were also numerous. In the metalimnion, these groups were replaced by scuticociliates and mixotrophic prostomatids. In the hypolimnion species known as benthic migrants appeared. We found a positive significant correlation (p < 0.05) between ciliate numbers and Chl a and bacterial densities. Only in the hypolimnion, the correlation between ciliates numbers and Chl a was not significant.     

Studies on ciliated protozoa in eutrophic lakes: 2. Field and laboratory studies on the effects of oxygen and other chemical gradients on ciliate distribution

An investigation into the spatial distribution of hypolimnetic ciliates in three small eutrophic lakes during the period of summer stratification was carried out. Peak ciliate densities were found to occur at the oxic/anoxic boundary, ciliate numbers declining with increasing depth within the hypolimnion. The ciliates only occurred in aerobic water where oxygen levels were less than about 0.5 mgl^–1 Laboratory experiments demonstrated that the ciliates swim upwards under anaerobic conditions but swim rapidly downwards under aerobic conditions. Further laboratory experiments showed that although the bulk of the population occured within anaerobic water, the hypolimnetic ciliates are aerobes and cannot survive indefinite anoxia. Despite the demonstrable toxicity of high levels of ammonia and sulphide, it was probably excesive distance from an available source of oxygen that excluded the ciliates from the lowest levels of the hypolimnion. Possible mechanisms which allowed these aerobic ciliates to colonise anaerobic water are considered.     

Analysis of the community structure of planktonic ciliated protozoa relative to trophic state in Florida lakes

The planktonic ciliate populations of 30 Florida lakes constituting a broad trophic gradient were examined to determine the response of protozoan community structure to increasing eutrophication. Both ciliate abundance and biomass were strongly related to lake trophic state. Comparison of the Florida data base with a comparable north temperate lake group indicated that subtropical lakes generally possess higher ciliate abundance and biomass at a given trophic state than temperate lakes. However the equations derived for each data base were not significantly different. Community diversity and species richness increased with increasing lake productivity. Highly acidic lakes displayed significantly reduced diversity and numbers of species when contrasted with nonacidic oligotrophic lakes. Small-bodied (< 30 um) ciliates dominated all lakes but were proportionally less important in oligotrophic lakes. Presence-absence data produced three assemblages: an ubiquitous association of primarily small ciliate taxa, a group of large ciliates mainly restricted to eutrophic-hypereutrophic lakes, and a very large ciliate,Stentor niger, which dominated the protozoan communities of acidic oligotrophic lakes.     

The temporal and spatial distribution of planktonic and benthic protozoan communities in a small productive lake

An investigation into the spatial and temporal distribution of protozoa in Esthwaite Water, a small eutrophic lake in the English Lake District, was carried out from mid-June until late September, 1977, during the period of summer stratification. Quantitative analyses of planktonic protozoan populations were performed on water samples collected at 1 m intervals throughout the water column and individual depth-time distribution profiles were constructed for the major ciliate species. Population density and species succession of the benthic protozoa was also studied.     

Phosphorus,Nitrogen and Chlorophyll-a Are Significant Factors Controlling Ciliate Communities in Summer in the Northern Beibu Gulf,South China Sea

Ciliates (protozoa) are ubiquitous components of plankton community and play important roles in aquatic ecosystems in regards of their abundance, biomass, diversity and energy turnover. Based on the stratified samples collected from the northern Beibu Gulf in August 2011, species composition, abundance, biomass, diversity and spatial pattern of planktonic ciliates were studied. Furthermore the main environmental factors controlling ciliate communities were determined. A total of 101 species belonging to 44 genera and 7 orders (i.e., Oligotrichida, Haptorida, Euplotida, Sessilida, Pleurostomatida, Scuticociliatida and Tintinnida) were identified. The variation of ciliate communities was significant at horizontal level, but that was not at vertical level. Based on cluster analysis, ciliate communities were divided into three main groups. Redundancy analysis (RDA) revealed that Group A, existing in the waters with higher concentration of phosphorus and nitrogen, was dominated by Tintinnidium primitivum. Group B in the waters with lower temperature and chlorophyll-a concentration, was dominated by Leegaardiella ovalis. Group C, existing in the waters with higher temperature and chlorophyll-a concentration, was dominated by large Strombidium spp. and Mesodinium rubrum. Combining multiple analytic methods, our results strongly supported that phosphorus, nitrogen and chlorophyll-a were the most significant factors affecting the ciliate communities in the northern Beibu Gulf in summer. Concentration of phosphorus and nitrogen primarily influenced ciliate biomass, implying a potential impact of eutrophication on ciliate growth. The correlation with chlorophyll-a concentration, on one hand indicate the response of ciliates to the food availability, and on the other hand, the ciliates containing chloroplasts or endosymbionts may contribute greatly to the chlorophyll-a.     

Ciliate communities of a large shallow lake: Association with macrophyte beds

We investigated the influence of macrophyte composition on ciliate community structure in a large, shallow, eutrophic Lake Võrtsjärv. We hypothesized that macrophyte composition must have strong influence on the dispersal of ecologically different ciliate groups in a shallow lake and that more diverse macrophyte stands cause also a greater diversity in the ciliate community. In Võrtsjärv macrophyte distribution is spatially strongly polarized both in east–west and north–south directions in relation to abiotic factors. Phragmites australis and Myriophyllum spicatum were the most widespread species occurring in most parts of the lake. Correlation of environmental, macrophyte and planktonic ciliate variables confirmed the suggested spatial gradients. More diverse macrophyte stands supported a high species richness and abundance of epiplanktonic community but showed negative influence on the number and abundance of euplanktonic ciliate taxa. Opposite trends were found relative to the abundance of P. australis. Benthic ciliates showed a similar distribution pattern to euplanktonic taxa being most abundant in sites were the Shannon–Weaver index for macrophytes was low. Strong polarizing effect of the lake's vegetation on planktonic ciliate diversity was reflected in correlations of the number of ciliate taxa as well as the numbers of eu- and epiplanktonic taxa with geographic co-ordinates.     

Seasonality of planktonic ciliated protozoa in 20 subtropical Florida lakes of varying trophic state

The planktonic ciliate populations of 20 Florida lakes ranging from oligotrophic to hypereutrophic were examined monthly for one year. Oligotrophic lakes displayed abundance peaks during fall mixis and biomass peaks in late winter and fall. Mesotrophic systems exhibited a spring-fall bimodality in ciliate abundance with a biomass maxima occurring during fall. Eutrophic/hypereutrophic lakes had pronounced abundance and biomass maxima during summer, with the large ciliates Plagiopyla nasuta and Paramecium trichium often contributing heavily to the midsummer biomass peak. Members of the Oligotrichida numerically dominated abundance and biomass peaks in oligotrophic lakes while the Scuticociliatida dominated the communities of higher trophic states. Total ciliate abundance and biomass were strongly correlated with chlorophyll a concentrations as were various ciliate taxonomic groups. The relationship between ciliate seasonal distribution in these subtropical lakes with lake thermal regimes and trophic state is discussed.     

Ciliate community structure and interactions within the planktonic food web in two alpine lakes of contrasting transparency

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A comparative study on planktonic ciliates in two shallow mesotrophic lakes (China): species composition, distribution and quantitative importance

The species richness and seasonal development of planktonic ciliates were studied and compared in two shallow mesotrophic lakes, one covered with dense submerged macrophytes, the other macrophyte poor. Considerable differences in ciliate species composition, dominant taxa, abundance and biomass were observed. Ciliates were much more species rich in the macrophyte-rich lake, while they were more abundant numerically in the macrophyte-poor lake. Altogether, 96 species, included in 53 genera, 14 orders were identified. Among them, 80 species (included in 45 genera, 14 orders) observed from the macrophyte-rich lake, against 49 species (36 genera, 12 orders) were from the macrophyte-poor lake. In the macrophyte-rich lake, the mean abundance and biomass were 13.5 cells ml^-1 and 547.10 g l^-1 f.w.; abundance and biomass were higher in spring and winter; naked oligotrichs dominated total ciliate abundance and Peritrichida dominated the biomass. In the macrophyte-poor lake, ciliate mean abundance and biomass were 35.5 cells ml^-1 and 953.39 g l^-1 f.w.; abundances peaked in autumn; Scuticociliates dominated the abundance and Tintinnids dominated the biomass. Possible causes for the observed differences are discussed.     

Protistan bacterivory in an oligomesotrophic lake: Importance of attached ciliates and flagellates

Seasonal and depth variations of the abundance, biomass, and bacterivory of protozoa (heterotrophic and mixotrophic flagellates and ciliates) were determined during thermal stratification in an oligomesotrophic lake (Lake Pavin, France). Maximal densities of heterotrophic flagellates (1.9 × 10³ cells ml^–1) and ciliates (6.1 cells ml^–1) were found in the metalimnion. Pigmented flagellates dominated the flagellate biomass in the euphotic zone. Community composition of ciliated protists varied greatly with depth, and both the abundance and biomass of ciliates was dominated by oligotrichs. Heterotrophic flagellates dominated grazing, accounting for 84% of total protistan bacterivory. Maximal grazing impact of heterotrophic flagellates was 18.9 × 10⁶ bacteria 1^–1h^–1. On average, 62% of nonpigmented flagellates were found to ingest particles. Ciliates and mixotrophic flagellates averaged 13% and 3% of protistan bacterivory, respectively. Attached protozoa (ciliates and flagellates) were found to colonize the diatom Asterionella formosa. Attached bacterivores had higher ingestion rates than free bacterivorous protozoa and may account for 66% of total protozoa bacterivory. Our results indicated that even in low numbers, epibiotic protozoa may have a major grazing impact on free bacteria. Correspondence: C. Amblard.     

The role of ciliated protozoa in pelagic freshwater ecosystems

The abundance and biomass of ciliates are both strongly related to lake trophic status as measured by chlorophylla concentrations. Taxonomic replacements occur with increasing eutrophication such that large-bodied forms (predominantly oligotrichs) are progressively replaced by smaller-bodied ciliates (mainly scuticociliates). Highly acidic lakes display a more pronounced dominance of large-bodied forms when contrasted with less acidic lakes of comparable trophy. Community structure of ciliate populations is determined largely by lake trophy with acidic oligotrophic systems being characterized by reduced diversity and species richness compared with hypereutrophic systems. The temporal and spatial distribution of small (< 100m) ciliate populations is ascribed to lake thermal regimes which provide localized concentrations of food resources. Likewise, in extremely productive lakes, very large (> 100m) meroplanktonic ciliates enter the water column during midsummer after the development of thermal stratification and associated profundal deoxygenation. Laboratory studies indicate that large zooplankton (crustaceans) are capable of utilizing ciliates as a food source, but there is little direct evidence from field studies documenting this trophic link. Ciliates can be voracious grazers of both bacterioplankton and phytoplankton, and each species has a distinct range of preferred particle size which is a function of both mouth size and morphology. Myxotrophic ciliates may be important components in some plankton communities, particularly during periods of nutrient limitation or after their displacement from the benthos of eutrophic lakes. Evidence regarding the importance of planktonic ciliated protozoa in nutrient regeneration and as intermediaries in energy flow is discussed.     

The Contribution of Ciliated Protozoa to Zooplankton Biomass in an Acidic, Subtropical Lake

Ciliated protozoa accounted for up to 50% of the mean daily zooplankton biomass in McCloud Lake, a small (5 ha), oligotrophic, acidic (pH 4.7) lake in north-central Florida. Food resources (algae and bacterioplankton) were limiting for crustacean and rotifer zooplankton during much of the year. Myxotrophic ciliates were a dominant component of the planktonic food web. Stentor niger , an uncommon species in the plankton of lakes, dominated the ciliate assemblage and usually comprised >90% of total ciliate biomass. Stentor niger always contained high densities of photosynthetic zoochlorellae and contributed an estimated 30% to the total autotrophic biomass.     

Fine scale spatial correlations between planktonic ciliates and dinoflagellates

Fine scale spatial distributions of planktonic ciliates whichprey on dinoflagellates were investigated in a small estuary.The horizontal distributions of Favella sp., Balanion sp. andStrobilidium sp. were positively correlated with the patchydistribution of dinoflagellates. The vertical distribution ofFavella was positively corrdated with the distribution of dinoflagellates.Throughout the diel cycle, the vertical distribution of Faveilawas similar to the vertical distribution of dinoflagellates.We speculate that behavioral responses are responsible for theassociation of diliates with their prey in the water column.Vertically coincident ciliate and algal populations should respondsimilarly to turbulent mixing and water displacement causedby wind stress, bathymetry, or frontal convergences. This wouldresult in horizontal patches in which the ciliates and algaeremain associated. This fine scale spatial coupling betweenciliates and their prey should result in higher ciliate growthrates and greater impact of ciliate grazing on phytoplanktonpopulations than would be predicted from average ciliate oralgal densities. Coincident patches of algae and ciliates mayalso provide higher food concentrations for larger grazers whichcan use both resources. ^{Contribution No. 5532, W.H.O.I.}     

The ciliated protozoa of the monomictic Lake Kinneret (Israel): species composition and distribution during stratification

The ciliate taxa from epilimnion, hypolimnion, and littoral of Lake Kinneret were studied from November to December 1987 during stratification. Two collections of planktonic and benthic ciliates were taken at each of six sampling stations. Benthic ciliates from the littoral zone were collected at seven stations along the lake shore. Densities were determined for each ciliate species and for total protozoans at each station and sampling date. Thirty-six species of ciliates, representing 31 Genera, 18 Orders and 7 Classes, were identified. Planktonic protozoans characteristic of the epilimnion included Coleps hirtus and Vorticella mayeri. The more abundant taxa of benthic ciliates in the profundal of the lake included Saprodinium dentatum, Plagiopyla nasuta and Dexiotricha plagia. The predominant ciliate in the littoral was Pleuronema coronatum. Principal component analysis, performed on the correlation matrix of both sampling stations and species, revealed that epilimnion, hypolimnion, and littoral belt were colonized by different ciliate communities.     

Size spectra and abundance of planktonic ciliates within various habitats in a macrophyte-dominated lake (Eastern Poland)

Body size, community structure, abundance and biomass of ciliates were compared in various stands of macrophytes in a macrophyte-abundant shallow lake in Eastern Poland. Samples were collected in belts of Phragmites, Typha, Ceratophyllum, Elodea, Stratiotes and Chara. Additionally, protozooplankton was collected from the open water zone surrounding the vegetation belts. Differences in numbers of ciliate taxa between micro-sites were statistically significant. The highest numbers were found in Chara and Ceratophyllum stands, lower numbers in Stratiotes and Elodea stands and the lowest in the open water, Phragmites and Typha areas. Ciliate biomass was, like density, significantly higher in submerged macrophytes than in emergent macrophytes and open water zones. Based on differences in macrophyte structure, two groups of habitats with similar patterns of size-related ciliate distribution were distinguished. The first group consisted of two vegetated zones of sparse stem structure (Phragmites and Typha) and the open water zone, the second group comprised submerged macrophyte species, which were more dense and complex. Generally, the abundance of ciliates correlated positively with total suspension solid (TSS) and total organic carbon (TOC) concentrations. In the Chara and Ceratophyllum stands, relations between ciliate numbers, TSS and TOC were stronger.     

Trophic relationships between bacteria and protozoa in the hypolimnion of a meromictic mesostrophic lake

The development of ciliated protozoan biomass in the hypolimnion of Piburger See, a small subalpine lake, was demonstrated to depend mainly on two factors. Firstly, the availability of oxygen or nitrate as electron acceptors determines the depth profiles of ciliates. Large quantities of ciliates and even maximum numbers were found at depths where no oxygen could be detected. If nitrate also disappeared during the summer stagnation period, the biomass of protozoa was strongly reduced. Nitrite peaks generally corresponded with ciliate peaks. An extension of Finlay's findings (dissimilatory nitrate reductase within the inner mitochondrial membrane) to other ciliate groups is hypothesized.Secondly, the biomass development of hypolimnetic ciliates was strongly correlated with the bacterial biomass registered approx. 2 weeks before (r² = 0.891, n = 14). The biomass of bacteria, on the other hand, was dependent upon the sedimentation rate of organic carbon (r² = 0.850, n = 15), if a time lag of approx. 2 weeks was taken into account. Therefore a total time lag of approx. 4 weeks was assumed to take place between sedimentation of organic substance and the corresponding increase in ciliate biomass (r² = 0.853, n = 14). Bacteria were shown to be an important intermediate link in the food chain of the hypolimnion. They appear to represent the principal energy source for pelagic ciliates. Sedimentation of organic carbon, nitrogen and phosphorus is the driving force for the establishment of the hypolimnetic microbial community.     

Contribution of ciliated protozoa to the planktonic biomass in a series of Ontario lakes: quantitative estimates and dynamical relationships

The plankton of nine Ontario lakes spanning several physiographicregions was sampled every two weeks during the ice-free periodof 1981, and one lake was studied in the three previous years.Phytoplankton, zooplankton, and ciliated protozoa were sampled,counted and sized. The size data were converted to biomass estimatesto yield quantitative comparisons of the relative allocationof biomass among different functional compartments. This isthe first study to look simultaneously and quantitatively atthe total plankton system of lakes (including ciliates, pbytoplanktonand net zooplankton) over a broad physiographic region. Ciliatesconstitute –10% of the non-algal biomass and 5% of thetotal planktonic biomass of these lakes. Ciliate standing cropsamong lakes are significantly corrrelated with total organicand total inorganic carbon concentrations in the water column,while the dynamics of ciliate biomass fluctuations are significantlycorrelated with variations in total phosphorus concentration,in conductivity, in Kjeldahl nitrogen concentration, and ininorganic carbon content. There appears to be a significantdynamical relationship between ciliates as a proportion of thetotal planktonic biomass, exclusive of filamentous and large(>30 µm) spherical algae, and the relative biomassof small algae (2–5 µm) as a fraction of total algalbiomass, again exclusive of filaments and large (>30 µm)algae. The hypothesis is advanced that ciliates primarily functionas bacterial grazers in planktonic ecosystems and that theirprimary competitors in this role are rotifers.     

Annual Changes of Abundance and Biomass of Planktonic Ciliates in the Gdańsk Basin,Southern Baltic

Seasonal changes in the species composition, abundance and biomass of planktonic ciliates were determined every 2–3 weeks at two sites of 30 m depth and one location of 105 m depth in the southwestern Gdańsk Basin between January 1987 and January 1988. A total of 40 ciliate taxa were observed during this period. Autotrophic Mesodinium rubrum dominated ciliate abundance and biomass: maximal values of 50 · 10⁻¹ ind. ^1-1 and 65 μg C 1⁻¹ were recorded. The annual mean biomass of M. rubrum comprised 6 to 9% of the annual mean phytoplankton biomass. The highest abundances and biomasses of heterotrophic ciliates were noted at all stations in the spring and summer in the euphotic zone with maximum values of 28 · 10³ ind. 1⁻¹ and 23 μg C 1⁻¹. Three ciliates assemblages were distinguished in the epipelagic layer: large and medium-size non-predatory ciliates, achieving peak abundance in spring and autumn; small-size microphagous ciliates and epibiotic ciliates which were abundant in summer, and large-size predacious ciliates dominating in spring. Below 60 m, a separate deep-water ciliate community composed of Prorodon-like ciliates and Metacystis spp. was found. The ciliate biomass in the 60–105 m layer was similar to the ciliate biomass in the euphotic zone. The heterotrophic ciliate community contributed 10 to 13% to the annual mean zooplankton biomass. The potential annual production of M. rubrum comprised 6 to 9% of the total primary production. Carbon demand of non-predatory ciliates, calculated on the basis of their potential production, was estimated to be equivalent to 12–15% of the gross primary production.