Phytoplankton and pelagic primary productivity in Øvre Heimdalsvatn

A phytoplankton investigation was carried out in the subalpine, low-productive Norwegian lake Øvre Heimdalsvatn in 1969–70 and 1972. This paper describes the temporal and spatial distribution of the standing stock of phytoplankton, and phytoplankton primary productivity. The annual average primary productivity in 1972 was 4.0–4.9 mg C m⁻³ d⁻¹; the annual average standing stock varied from 120 mg m⁻³ (freshweight) in 1969–70, to 250 mg m⁻³ in 1972. Phytoplankton species composition and size distribution is discussed. Throughout the year the phytoplankton is dominated by small (ultraplankton) species; μ-algae (< 5 μm) showed cell concentrations up to 15 mill. cells 1⁻¹. The dominating group was chrysophytes; cryptophytes, dinoflagellates or green algae were at times abundant. A phytoplankton monthly budget and a diagram showing annual average carbon flow through the standing stock of phytoplankton are presented; the phytoplankton dynamics in Øvre Heimdalsvatn is compared to that of other low-productive lakes.     

Periphytic and drifting microalgae in a tributary stream of Øvre Heimdalsvatn

Samples of drifting and periphytic microalgae were collected during 1972 from a fast-flowing, stony stream (Brurskardsbekken) in the Jotunheimen mountain area, central southern Norway. The predominant algal groups in the drift and the periphyton were diatoms and green algae, while only a few species were recorded in both communities. A considerable number of species from Chrysophyceae, Cryptophyceae and other algal classes were also recorded in the drift samples. The species composition was in good agreement with microalgal communities earlier described from mountain areas in Scandinavia, although species which probably are new to Norway, were also recorded. A quantitatively important fraction of planktonic species in the drift is interpreted as a contribution from lacustrine habitats in the watercourse. A general change in the periphyton during the summer, from green algae to diatoms, was observed, Altitudinal differences in the periphyton included a delayed green algal maximum at higher altitudes compared to lower. In the zone around the upper birch limit, a transition in species composition as one goes up stream, described in other investigations, was not observed.     

Quantitative responses of lake phytoplankton to eutrophication in Northern Europe

Based on the currently largest available dataset of phytoplankton in lakes in northern Europe, we quantified the responses of three major phytoplankton classes to eutrophication. Responses were quantified by modelling the proportional biovolumes of a given group along the eutrophication gradient, using generalized additive models. Chlorophyll-a (Chl-a) was chosen as a proxy for eutrophication because all classes showed more consistent responses to Chl-a than to total phosphorus. Chrysophytes often dominate in (ultra-) oligotrophic lakes, and showed a clear decrease along the eutrophication gradient. Pennate diatoms were found to be most abundant at moderate eutrophication level (spring-samples). Cyanobacteria often dominate under eutrophic conditions, especially in clearwater lakes at Chl-a levels >10 μg l⁻¹ (late summer samples). We compare the relationships among types of lakes, based on the lake typology of the northern geographic intercalibration group, and among countries sharing common lake types. Significant differences were found especially between humic and clearwater lakes, and between low- and moderately alkaline lakes, but we could not identify significant differences between shallow and deep lakes. Country-specific differences in response curves were especially pronounced between lakes in Norway and Finland, while Swedish lakes showed an intermediate pattern, indicating that country-specific differences reflect large-scale geographic and climatic differences in the study area.     

Nutrient Enrichment and Planktonic Biomass Ratios in Lakes

Phosphorus (P) to chlorophyll ratios and zooplankton–phytoplankton (Z:P) biomass ratios were assessed in 400 temperate lakes over a gradient of phosphorus (P) and with different fish communities. Most of the lakes in this survey were oligotrophic, with a median total P of 7.3 μg P L⁻¹. Thus, the survey provided information on food web effects during the early phase of eutrophication. There was no tendency toward a reduced yield of autotrophs per unit of P over the gradient covered in this survey. The zooplankton yield per unit of P or chlorophyll a decreased slightly with increased nutrient concentrations, and Z:P biomass ratios decreased with fish community classes, reflecting increased fish predation pressure. However, the variability in biomass ratios within a given range of P and fish class was some 100 times higher than the difference over the gradients. This finding suggests that lake-specific properties, community composition, and food quality are by far the most important determinants of biomass ratios and probably also trophic efficiency in lakes; it further suggests that these factors are superimposed on the general effect of eutrophication, at least up to 30 μg P L⁻¹.     

Regional species pools control community saturation in lake phytoplankton

Recent research has highlighted that positive biodiversity–ecosystem functioning relationships hold for all groups of organisms, including microbes. Yet, we still lack understanding regarding the drivers of microbial diversity, in particular, whether diversity of microbial communities is a matter of local factors, or whether metacommunities are of similar importance to what is known from higher organisms. Here, we explore the driving forces behind spatial variability in lake phytoplankton diversity in Fennoscandia. While phytoplankton biovolume is best predicted by local phosphorus concentrations, phytoplankton diversity (measured as genus richness, G) only showed weak correlations with local concentrations of total phosphorus. By estimating spatial averages of total phosphorus concentrations on various scales from an independent, spatially representative lake survey, we found that close to 70 per cent of the variability in local phytoplankton diversity can be explained by regionally averaged phosphorus concentrations on a scale between 100 and 400 km. Thus, the data strongly indicate the existence of metacommunities on this scale. Furthermore, we show a strong dependency between lake productivity and spatial community turnover. Thus, regional productivity affects beta-diversity by controlling spatial community turnover, resulting in scale-dependent productivity-diversity relationships. As an illustration of the interaction between local and regional processes in shaping microbial diversity, our results offer both empirical support and a plausible mechanism for the existence of common scaling rules in both the macrobial and the microbial worlds. We argue that awareness of regional species pools in phytoplankton and other unicellular organisms may critically improve our understanding of ecosystems and their susceptibility to anthropogenic stressors.     

The phytoplankton of Lake Atnsjøen, Norway – a long-term investigation

Quantitative samples were collected from Lake Atnsjøen five times per year in the growth seasons 1990–2000. The samples were analysed for variation in the phytoplankton composition, and the total volume and volume of the main groups of algae were calculated. Lake Atnsjøen is a large, deep and unregulated lake with a surface area of 4.8 km² and a maximum depth of 80.2 m. It is a nutrient-poor, oligotrophic lake with a maximum phytoplankton volume varying between 125–393 mm³/m³ in the years 1990–2000. The phytoplankton community is dominated by species of the groups Chrysophyceae and Cryptophyceae. The chrysophytes dominate the phytoplankton in the early part of the growth season (May–June) while the cryptophytes increase throughout the season and dominate in the autumn. Among the chrysophytes different species of chrysomonads were most frequent together with common species of the genus Dinobryon like D. borgei, D. cylindricum var. alpinum and D. crenulatum. A total of 22 species or taxa of chrysophytes were recorded in the samples. Common among the cryptomonads were several species of the genus Cryptomonas. Most important quantitatively, however, were Rhodomonas lacustris and Katablepharis ovalis. The succesion of the phytoplankton throughout the growth season was similar from year to year in quantitative as well as qualitative terms, but some changes were recorded after the great flood in 1995. Canonical Correspondence Analysis (CCA) shows a slight, but significant, phytoplankton community change over the succeeding years.     

Performance of a new phytoplankton composition metric along a eutrophication gradient in Nordic lakes

A new phytoplankton metric is presented, which is developed from a large dataset of Norwegian lakes (>2,000 samples from >400 lakes). In contrast to previous metrics, this index is not built on selected ‘indicative’ taxa, but uses all available taxonomic information at genus and species level. Taxa optima with respect to lake trophic status (derived from total phosphorus concentrations) are used to calculate a phytoplankton trophic index (TI) for each sample. Analysis of the TI shows that phytoplankton communities exhibit highly non-linear responses to eutrophication in Norwegian lakes. Reference lakes are characterized by very similar TIs despite having considerable variation in total phosphorus and chlorophyll a concentrations. TI exhibits a non-linear distribution along the eutrophication gradient which separates unimpacted from impacted sites in the study area. We further show that TI exhibits smaller seasonal variations than chlorophyll a, making it a more reliable indicator for lake monitoring. Implications for its applicability within the WFD are discussed.     

Autotroph:herbivore biomass ratios; carbon deficits judged from plankton data

A survey on phytoplankton:zooplankton biomass ratios was performed in 342 Norwegian lakes, covering a wide range in lake size and productivity (total phosphorus: 3–246 g l^–1), but with most localities being oligo- to mesotrophic. Mean phytoplankton biomass was 88 g C l^–1, yet with the majority below 50 g C l^–1and a median of 25 g C l^–1. Total zooplankton biomass displayed a mean and median of 37 and 26 g C l^–1, respectively. Cladocerans were by far the dominant group, making up a median of almost 60% of total zooplankton biomass. Total zooplankton biomass as well as that of major aggregated metazoan taxa (cladocerans, calanoid copepods, cyclopoid copepods and rotifers) all showed a positive, but weak correlation with total phytoplankton biomass. These weak correlations suggest that algal biomass per se is a poor predictor of zooplankton biomass. An average phyto-:zooplankton biomass ratio (C:C) of 2.8 (SD±4.7) was found. 30% of the lakes had a phyto-:zooplankton biomass ratio below unity. While there was no correlation between the phyto-:zooplankton biomass ratio with increasing productivity in terms of P concentration, there was a higher biomass ratio in lakes with high fish predation pressure. The low ratio of phyto-:zooplankton biomass suggest major requirements from non-algal sources of C in the zooplankton diet. The need for dietary subsidizing is also supported by the fact that more than 75% of the lakes had algal biomass less than the estimated threshold for net positive growth of zooplankton, although it should be kept in mind that a high share of picoplankton would imply an underestimation of autotroph biomass in these lakes. Since the C-deficiency apparently is most pronounced in oligotrophic systems, it contradicts the view that the detritus pathways plays a predominant role in highly productive systems only, but while the source of detritus probably is mostly of autochthonous origin in eutrophic lakes, allochthonous detritus will be more important in oligotrophic systems.     

The Impact of Nutrient State and Lake Depth on Top-down Control in the Pelagic Zone of Lakes: A Study of 466 Lakes from the Temperate Zone to the Arctic

Using empirical data from 466 temperate to arctic lakes covering a total phosphorus (TP) gradient of 2-1036 mg L-1, we describe how the relative contributions of resource supply, and predator control change along a nutrient gradient. We argue that (a) predator control on large-bodied zooplankton is unimodally related to TP and is highest in the most nutrient-rich and nutrient-poor lakes and generally higher in shallow than deep lakes, (b) the cascading effect of changes in predator control on phytoplankton decreases with increasing TP, and (c) these general patterns occur with significant variations--that is, the predation pressure can be low or high at all nutrient levels. A quantile regression revealed that the median share of the predator-sensitive Daphnia to the total cladoceran biomass was significantly related unimodally to TP, while the 10% and 90% percentiles approached 0 and 100%, respectively, at all TP levels. Moreover, deep lakes (more than 6 m) had a higher percentage of Daphnia than shallow (less than 6 m) lakes. The median percentage of Daphnia peaked at 0.15 mg L-1 in shallow lakes and 0.09 mg L-1 in deep lakes. The assumption that fish are responsible for the unimodality was supported by data on the abundance of potential planktivorous fish (catch net-1 night-1 gill nets with the different mesh sizes [CPUE]). To elucidate the potential cascading effect on phytoplankton, we examined the zooplankton phytoplankton biomass ratio. Even though this ratio was inversely related to CPUE at all TP levels, we found an overall higher ratio in oligotrophic lakes that declined toward low values (typically below 0.2) in hypertrophic lakes. These results suggest that planktivorous fish have a more limited effect on the grazing control of phytoplankton in oligotrophic lakes than in eutrophic lakes, despite similar predator control of large-bodied zooplankton. Accordingly, the phytoplankton yield, expressed as the chlorophyll a-TP ratio, did not relate to CPUE at low TP, but it increased significantly with CPUE at high TP. We conclude that the chances of implementing a successful restoration program using biomanipulation as a tool to reduce phytoplankton biomass increase progressively with increasing TP, but that success in the long term is most likely achieved at intermediate TP concentrations.     

An annotated check-list of microalgae from Øvre Heimdalsvatn and its tributaries

About one hundred genera with more than 300 species have been identified in water samples, net haul material, periphyton samples from artificial substrate, and benthos and sediment samples collected during 1969–1972. More than 40 taxa of lower rank (subspecies, varieties, forms) were recorded. The most prominent algal groups were diatoms with 138 taxa, and green algae with 122 taxa of which the majority was desmids. The material from the epipelic and epilithic communities in the lake is very scarce, while the list of plankton species is relatively complete, with the exception that a large number of small flagellates could not be identified to species level. The taxa which were identified, are presented in a list with additional notes on occurrence.