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Phytoplankton dynamics in a shallow eutrophic lake were investigated over a 3-year period with respect to environmental forces which drive species composition and diversity. Diversity was calculated on the basis of species as well as on the basis of their functional properties (the C-R-S-concept). Stratification and water column mixing had a strong impact on phytoplankton composition. Application of a similarity-diversity model revealed that a high diversity was a transient non-stable state, whereas drastic changes or long-lasting stable environmental conditions are characterized by low diversity. This effect was more pronounced when the diversity was calculated on the basis of the phytoplankton species functional properties. Thus, this functional approach supports the intermediate disturbance hypothesis from field data.
The mode of nutrition of mixotrophic flagellates determines the food quality for their consumers
(2007)
The invasion success of the cyanobacterium Cylindrospermopsis raciborskii in experimental mesocosms
(2017)
The potentially toxic, invasive cyanobacterium Cylindrospermopsis raciborskii, originating from sub-tropical regions, has spread into temperate climate zones in almost all continents. Potential factors in its success are temperature, light and nutrient levels. Grazing losses through zooplankton have been measured in the laboratory but are typically not regarded as a factor in (failed) invasion success. In some potentially suitable lakes, C. raciborskii has never been found, although it is present in water bodies close by. Therefore, we tested the invasive potential of three different isolates introduced into natural plankton communities using laboratory mesocosm experiments under three grazing levels: ambient zooplankton densities, removal of large species using 100 mu m mesh and a ca. doubling of large species. Three C. raciborskii isolates originating from the same geographic region (North-East Germany) were added separately to the four replicates of each treatment and kept in semi-continuous cultures for 21 days. Two isolates disappeared from the mesocosms and were also not viable in filtered lake water indicating that the lake water itself or the switch from culture medium to lake water led to the decay of the inoculated C. raciborskii. Only one out of the three isolates persisted in the plankton communities at a rather low level and only in the treatment without larger zooplankton. This result demonstrates that under potentially suitable environmental conditions, top-down control from zooplankton might hamper the establishment of C. raciborskii. Non-metric multidimensional scaling showed distinct variation in resident phytoplankton communities between the different grazing levels, thus differential grazing impact shaped the resident community in different ways allowing C. raciborskii only to invade under competitive (= low grazing pressure) conditions. Furthermore, even after invasion failure, the temporary presence of C. raciborskii influenced the phytoplankton community.
In most biodiversity studies, taxonomic diversity is the measure for the multiplicity of species and is often considered to represent functional diversity. However, trends in taxonomic diversity and functional diversity may differ, for example, when many functionally similar but taxonomically different species co-occur in a community. The differences between these diversity measures are of particular interest in diversity research for understanding diversity patterns and their underlying mechanisms. We analysed a temporally highly resolved 20-year time series of lake phytoplankton to determine whether taxonomic diversity and functional diversity exhibit similar or contrasting seasonal patterns. We also calculated the functional mean of the community in n-dimensional trait space for each sampling day to gain further insights into the seasonal dynamics of the functional properties of the community. We found an overall weak positive relationship between taxonomic diversity and functional diversity with a distinct seasonal pattern. The two diversity measures showed synchronous behaviour from early spring to mid-summer and a more complex and diverging relationship from autumn to late winter. The functional mean of the community exhibited a recurrent annual pattern with the most prominent changes before and after the clear-water phase. From late autumn to winter, the functional mean of the community and functional diversity were relatively constant while taxonomic diversity declined, suggesting competitive exclusion during this period. A further decline in taxonomic diversity concomitant with increasing functional diversity in late winter to early spring is seen as a result of niche diversification together with competitive exclusion. Under these conditions, several different sets of traits are suitable to thrive, but within one set of functional traits only one, or very few, morphotypes can persist. Taxonomic diversity alone is a weak descriptor of trait diversity in phytoplankton. However, the combined analysis of taxonomic diversity and functional diversity, along with the functional mean of the community, allows for deeper insights into temporal patterns of community assembly and niche diversification.
Isolated extreme habitats are ideally suited to investigate pivotal ecological processes such as niche use, local adaptation and dispersal. Extremophilic animals living in isolated habitats face the problem that dispersal is limited through the absence of suitable dispersal corridors, which in turn facilitates local adaptation. We used five rotifer isolates from extremely acidic mining lakes with a pH of below 3 as model organisms to test whether these isolates are acidotolerant or acidophilic, whether they survive and reproduce at their niche edges (here pH 2 and circum-neutral pH) and whether local adaptation has evolved. To evaluate potential dispersal limitation, we tested whether animals and their parthenogenetic eggs survive and remain reproductive or viable at unfavourable pH-conditions. All five isolates were acidophilic with a pH-optimum in the range of 4-6, which is well above the pH (< 3) of their lakes of origin. At unfavourable high pH, in four out of the five isolates parthenogenetic females produced a high number of non-viable eggs. Females and eggs produced at favourable pH (4) remained vital at an otherwise unfavourable pH of 7, indicating that for dispersal no acidic dispersal corridors are necessary. Common garden experiments revealed no clear evidence for local adaptation in any of the five isolates. Despite their acidophilic nature, all five isolates can potentially disperse via circum-neutral water bodies as long as their residence time is short, suggesting a broader dispersal niche than their realized niche. Local adaptation might have been hampered by the low population sizes of the rotifers in their isolated habitat and the short time span the mining lakes have existed.
Lake morphometry and wind exposure may shape the plankton community structure in acidic mining lakes
(2010)
Acidic mining lakes (pH <3) are specific habitats exhibiting particular chemical and biological characteristics. The species richness is low and mixotrophy and omnivory are common features of the plankton food web in such lakes. The plankton community structure of mining lakes of different morphometry and mixing type but similar chemical characteristics (Lake 130, Germany and Lake Langau, Austria) was investigated. The focus was laid on the species composition, the trophic relationship between the phago-mixotrophic flagellate Ochromonas sp. and bacteria and the formation of a deep chlorophyll maximum along a vertical pH-gradient. The shallow wind-exposed Lake 130 exhibited a higher species richness than Lake Langau. This increase in species richness was made up mainly by mero-planktic species, suggesting a strong benthic/littoral - pelagic coupling. Based on the field data from both lakes, a nonlinear, negative relation between bacteria and Ochromonas biomass was found, suggesting that at an Ochromonas biomass below 50 mu g CL-1. the grazing pressure on bacteria is low and with increasing Ochromonas biomass bacteria decline. Furthermore, in Lake Langau, a prominent deep chlorophyll maximum was found with chlorophyll concentrations ca. 50 times higher than in the epilimnion which was build up by the euglenophyte Lepocinclis sp. We conclude that lake morphometry, and specific abiotic characteristics such as mixing behaviour influence the community structure in these mining lakes.
Water column mixing is known to have a decisive impact on plankton communities. The underlying mechanisms depend on the size and depth of the water body, the nutrient status, and the plankton community structure and they are well understood for shallow polymictic and deep stratified lakes. Two consecutive mixing events of similar intensity under different levels of herbivory were performed in enclosures in a shallow, but periodically stratified, eutrophic lake, in order to investigate the effects of water column mixing on bacteria abundance, phytoplankton abundance and diversity, and rotifer abundance and fecundity. When herbivory by filter-feeding zooplankton was low, water column mixing provoking a substantial nutrient input into the euphotic zone, led to an intense net increase of bacteria and phytoplankton biomass. Phytoplankton diversity was lower in the mixed enclosures than in the undisturbed ones owing to the larger contribution of a few fast-growing species. After the second mixing event at high biomass of filter-feeding crustaceans, the increase of phytoplankton biomass was lower than after the first mixing, and diversity remained unchanged as the enhanced growth of small fast-growing was prevented by zooplankton grazing. Bacteria abundance did not increase after the second mixing, when cladoceran biomass was high. Rotifer fecundity indicated a transmission of the phytoplankton response to the next trophic level. Our results suggest that water column mixing in shallow eutrophic lakes with periodic stratification has a strong effect on the plankton community by enhanced nutrient availability rather than resuspension or reduced light availability. This fuels the basis of the classic and microbial food chain via enhanced phytoplankton and bacteria growth, but the effects on biomass may be dampened by high levels of herbivory.
Planktische Räuber-Beute-Systeme : experimentelle Untersuchungen von ökologischen Synchronisationen
(2000)
Trait-based approaches have become increasingly successful in community ecology. They assume that the distribution of functional traits within communities responds in a predictable way to alterations in environmental forcing and that strong forcing may accelerate such trait changes. We used high frequency measurements of phytoplankton to test these assumptions. We analyzed the seasonal and long-term dynamics of the community trait mean within a multi-dimensional trait space under alternating multifactorial environmental conditions. The community trait mean exhibited a distinct recurrent annual pattern that reflected minor changes in climate, herbivory and nutrients. Independent of early spring conditions, the community trait mean was repeatedly driven into a narrow confined area in the trait space under pronounced herbivory during the clear water phase. The speed of movement was highest at the onset and the relaxation of such strong unidirectional forcing. Thus, our data support the conceptual framework of trait-based ecology that alterations in environmental conditions are systematically tracked by adjustments in the dominant functional trait values and that the speed of trait changes depends on the kind and intensity of the selection pressure. Our approach provides a sensitive tool to detect small functional differences in the community related to subtle differences in forcing.
Extreme habitats often harbor specific communities that differ substantially from non-extreme habitats. In many cases, these communities are characterized by archaea, bacteria and protists, whereas the number of species of metazoa and higher plants is relatively low. In extremely acidic habitats, mostly prokaryotes and protists thrive, and only very few metazoa thrive, for example, rotifers. Since many studies have investigated the physiology and ecology of individual species, there is still a gap in research on direct, trophic interactions among extremophiles. To fill this gap, we experimentally studied the trophic interactions between a predatory protist (Actinophrys sol, Heliozoa) and its prey, the rotifers Elosa woralli and Cephalodella sp., the ciliate Urosomoida sp. and the mixotrophic protist Chlamydomonas acidophila (a green phytoflagellate, Chlorophyta). We found substantial predation pressure on all animal prey. High densities of Chlamydomonas acidophila reduced the predation impact on the rotifers by interfering with the feeding behaviour of A. sol. These trophic relations represent a natural case of intraguild predation, with Chlamydomonas acidophila being the common prey and the rotifers/ciliate and A. sol being the intraguild prey and predator, respectively. We further studied this intraguild predation along a resource gradient using Cephalodella sp. as the intraguild prey. The interactions among the three species led to an increase in relative rotifer abundance with increasing resource (Chlamydomonas) densities. By applying a series of laboratory experiments, we revealed the complexity of trophic interactions within a natural extremophilic community.
Extreme habitats often harbor specific communities that differ substantially from non-extreme habitats. In many cases, these communities are characterized by archaea, bacteria and protists, whereas the number of species of metazoa and higher plants is relatively low. In extremely acidic habitats, mostly prokaryotes and protists thrive, and only very few metazoa thrive, for example, rotifers. Since many studies have investigated the physiology and ecology of individual species, there is still a gap in research on direct, trophic interactions among extremophiles. To fill this gap, we experimentally studied the trophic interactions between a predatory protist (Actinophrys sol, Heliozoa) and its prey, the rotifers Elosa woralli and Cephalodella sp., the ciliate Urosomoida sp. and the mixotrophic protist Chlamydomonas acidophila (a green phytoflagellate, Chlorophyta). We found substantial predation pressure on all animal prey. High densities of Chlamydomonas acidophila reduced the predation impact on the rotifers by interfering with the feeding behaviour of A. sol. These trophic relations represent a natural case of intraguild predation, with Chlamydomonas acidophila being the common prey and the rotifers/ciliate and A. sol being the intraguild prey and predator, respectively. We further studied this intraguild predation along a resource gradient using Cephalodella sp. as the intraguild prey. The interactions among the three species led to an increase in relative rotifer abundance with increasing resource (Chlamydomonas) densities. By applying a series of laboratory experiments, we revealed the complexity of trophic interactions within a natural extremophilic community.
Trait-based approaches to investigate (short- and long-term) phytoplankton dynamics and community assembly have become increasingly popular in freshwater and marine science. Although the nature of the pelagic habitat and the main phytoplankton taxa and ecology are relatively similar in both marine and freshwater systems, the lines of research have evolved, at least in part, separately. We compare and contrast the approaches adopted in marine and freshwater ecosystems with respect to phytoplankton functional traits. We note differences in study goals relating to functional trait use that assess community assembly and those that relate to ecosystem processes and biogeochemical cycling that affect the type of characteristics assigned as traits to phytoplankton taxa. Specific phytoplankton traits relevant for ecological function are examined in relation to
herbivory, amplitude of environmental change and spatial and temporal scales of study. Major differences are identified, including the shorter time scale for regular environmental change in freshwater ecosystems compared to that in the open oceans as well as the
type of sampling done by researchers based on site-accessibility. Overall, we encourage researchers to better motivate why they apply trait-based analyses to their studies and to make use of process-driven approaches, which are more common in marine studies. We further propose fully comparative trait studies conducted along the habitat gradient spanning freshwater to brackish to marine systems, or along geographic gradients. Such studies will benefit from the combined strength of both fields.
Trait-based approaches to investigate (short- and long-term) phytoplankton dynamics and community assembly have become increasingly popular in freshwater and marine science. Although the nature of the pelagic habitat and the main phytoplankton taxa and ecology are relatively similar in both marine and freshwater systems, the lines of research have evolved, at least in part, separately. We compare and contrast the approaches adopted in marine and freshwater ecosystems with respect to phytoplankton functional traits. We note differences in study goals relating to functional trait use that assess community assembly and those that relate to ecosystem processes and biogeochemical cycling that affect the type of characteristics assigned as traits to phytoplankton taxa. Specific phytoplankton traits relevant for ecological function are examined in relation to
herbivory, amplitude of environmental change and spatial and temporal scales of study. Major differences are identified, including the shorter time scale for regular environmental change in freshwater ecosystems compared to that in the open oceans as well as the
type of sampling done by researchers based on site-accessibility. Overall, we encourage researchers to better motivate why they apply trait-based analyses to their studies and to make use of process-driven approaches, which are more common in marine studies. We further propose fully comparative trait studies conducted along the habitat gradient spanning freshwater to brackish to marine systems, or along geographic gradients. Such studies will benefit from the combined strength of both fields.
1. The biovolume-specific carbon content, relative egg volume (a measure of per-offspring reproductive investment), growth and grazing rates, and the gross growth efficiency (GGE) of the rotifer Cephalodella hoodi, isolated from an extremely acidic habitat (pH 2.65), were determined and compared with literature values for rotifers living in circum-neutral habitats in order to reveal potential special features or adaptations related to the extreme habitat of C. hoodi. 2. Of the two dominant phytoflagellates (Ochromonas sp. and Chlamydomonas acidophila) that occur in the natural habitat of C. hoodi, only C. acidophila promoted positive growth and reproduction and, thus, the following results were obtained with C. acidophila as a food alga. 3. The body volume-specific carbon content of C. hoodi is in the range of that found in rotifers from circum-neutral lakes, suggesting that no costly carbon investment, brought about by the thickening of the lorica, for example, was required to withstand low pH. 4. The egg volume of C. hoodi exhibited no phenotypic plasticity dependent on the food concentration and, thus, C. hoodi allocated a constant, absolute amount of energy to each individual offspring. No adaptation to low food densities was found. 5. A dome-shaped type II functional response curve was found to describe the ingestion of Chlamydomonas as a source of food. 6. Compared with other rotifers, C. hoodi had a high threshold and half-saturating food concentration (=low affinity) but also a high maximum growth rate and a relatively high GGE, suggesting no severe adverse effect of low pH
Herbivore populations are commonly restricted by resource limitation, by predation or a combination of the two. Food supplement experiments are suitable for investigating the extent of food limitation at any given time. The main part of this study was performed in an extremely acidic lake (pH 2.7) where the food web consists of only a few components and potential food sources for herbivores are restricted to two flagellates. Life table experiments proved that Chlamydomonas was a suitable food source whereas Ochromonas was an unsuitable food source. The two flagellates and the two rotifers exhibit a pronounced vertical distribution pattern. In this study, a series of food supplement experiments were performed in order to: (1) quantify and compare potential resource limitation of two primary consumers (Cephalodella hoodi and Elosa worallii, Rotatoria) over time, (2) compare their response at different temperatures, (3) evaluate the effect of having an unsuitable food source alongside a valuable one, (4) estimate the effect of predation on rotifers by Heliozoa, and (5) compare the results with those from other acidic lakes. Additionally, the spatio- temporal population dynamics of both species were observed. The field data confirmed a vertical separation of the two species with E. worallii dominating in the upper water layers, and C. hoodi in the deeper, cooler water layers. The results from the food supplement experiments in which Chlamydomonas served as the supplemented suitable food source showed that the two rotifers were food limited in the epilimnion throughout the season to different extents, with Cephalodella being more severely food limited than Elosa. The experiments at different temperatures provided evidence that Elosa had a higher optimum temperature for growth than Cephalodella. When the unsuitable food algae Ochromonas was added alongside the suitable food source Chlamydomonas, C. hoodi was unaffected but E. worallii was negatively affected. Predation of Heliozoa on rotifers was observed but the total effect on the rotifer dynamics is probably low. The comparison with other lakes showed that resource limitation also occurred in one other lake, although to a lesser extent. Overall, the vertical separation of the two rotifers could be explained by both their differential extent of resource limitation and differential response to temperature.
1. This is a discussion of the applicability to the phytoplankton of the concepts of 'Plant Functional Types' (PFTs) and 'Functional Diversity' (FD), which originated in terrestrial plant ecology. 2. Functional traits driving the performance of phytoplankton species reflect important processes such as growth, sedimentation, grazing losses and nutrient acquisition. 3. This paper presents an objective, mathematical way of assigning PFTs and measuring FD. Ecologists can use this new approach to investigate general hypotheses (e.g. the intermediate disturbance hypothesis (IDH), the insurance hypothesis and synchronicity phenomena), since, for example, in its original formulation the IDH makes its predictions based on FD rather than species diversity.
According to resource allocation theory, animals face a trade off between the allocation of resources into reproduction and into individual growth/maintenance. This trade off is reinforced when food conditions decline. It is well established in biological research that many animals increase their life span when food is in suboptimal supply for growth and/or reproduction. Such a situation of reduced food availability is called dietary restriction. An increase in life span under dietary restricted conditions is seen as a strategy to tolerate periods of food shortage so that the animals can start reproduction again when food is in greater supply. In this study, the effect of dietary restriction on life span and reproduction in two rotifer species, Cephalodella sp. and Elosa worallii, was investigated using life table experiments. The food concentration under dietary restricted conditions was below the threshold for population growth. It was (1) tested whether the rotifers start reproduction again after food replenishment, and (2) estimated whether the time scale of dietary restricted conditions is relevant for the persistence of a population in the field. Only E. worallii responded to dietary restriction with an increase in life span at the expense of reproduction. After replenishment of food, E. worallii started to reproduce again within I day. With an increase in the duration of dietary restricted conditions of up to 15 days, which is longer than the median life span of E. worallii under food saturation, the life span increased and the life time reproduction decreased. These results suggest that in a temporally (or spatially) variable environment, some rotifer populations can persist even during long periods of severe food deprivation.
We investigated the morphology, phylogeny of the 18S rDNA, and pH response of Oxytricha acidotolerans sp. nov. and Urosomoida sp. (Ciliophora, Hypotricha) isolated from two chemically similar acid mining lakes (pH similar to 2.6) located at Langau, Austria, and in Lusatia, Germany. Oxytricha acidotolerans sp. nov. from Langau has 18 frontal-ventral-transverse cirri but a very indistinct kinety 3 fragmentation so that the assignment to Oxytricha is uncertain. The somewhat smaller species from Lusatia has a highly variable cirral pattern and the dorsal kineties arranged in the Urosomoida pattern and is, therefore, preliminary designated as Urosomoida sp. The pH response was measured as ciliate growth rates in laboratory experiments at pH ranging from 2.5 to 7.0. Our hypothesis was that the shape of the pH reaction norm would not differ between these closely related (3% difference in their SSU rDNA) species. Results revealed a broad pH niche for O. acidotolerans, with growth rates peaking at moderately acidic conditions (pH 5.2). Cyst formation was positively and linearly related to pH. Urosomoida sp. was more sensitive to pH and did not survive at circumneutral pH. Accordingly, we reject our hypothesis that similar habitats would harbour ciliate species with virtually identical pH reaction norm.