Abiotic drivers of interannual phytoplankton variability and a 1999–2000 regime shift in the North Sea examined by multivariate statistics

The Dutch coastal zone is a region of the North Sea with a marked interannual and long‐term abiotic and phytoplankton variability. To investigate the relationship between abiotic variability and phytoplankton composition, two routine water monitoring data sets (1991–2005) were examined. Multivariate statistics revealed two significant partitions in the data. The first consisted of interannual abiotic fluctuations that were correlated to Rhine discharge that affected the abundance of summer and autumn diatom species. The second partition was caused by a shift in the abiotic data from 1998 to 1999 that was followed by a shift in phytoplankton composition from 1999 to 2000. Important factors in the abiotic shift were decreases in suspended matter (SPM) and phosphate (DIP) concentrations, as well as in pH. The decrease in SPM was caused by a reduction in wind speed. The increase in water column daily irradiance from the decrease in SPM led to increases in the abundance of winter–spring species, notably the prymnesiophyte Phaeocystis globosa. Because wind speed is related to the North Atlantic Oscillation (NAO) index it was possible to correlate NAO index and P. globosa abundance. Only five abiotic variables representing interannual and long‐term variability, including Rhine discharge and NAO index, were needed to model the observed partitions in phytoplankton composition. It was concluded that interannual variability in the coastal phytoplankton composition was related to year‐to‐year changes in river discharge while the long‐term shift was caused by an alternating large‐scale meteorological phenomenon.     

Relationships between picophytoplankton and environmental variables in lakes along a gradient of water colour and nutrient content

SUMMARY 1. Biomass and production of picophytoplankton, phytoplankton and heterotrophic bacterioplankton were measured in seven lakes, exhibiting a broad range in water colour because of humic substances. The aim of the study was to identify environmental variables explaining the absolute and relative importance of picophytoplankton. In addition, two dystrophic lakes were fertilised with inorganic phosphorus and nitrogen, to test eventual nutrient limitation of picophytoplankton in these systems.
2. Picophytoplankton biomass and production were highest in lakes with low concentrations of dissolved organic carbon (DOC), and DOC proved the factor explaining most variation in picophytoplankton biomass and production. The relationship between picophytoplankton and lake trophy was negative, most likely because much P was bound in humic complexes. Picophytoplankton biomass decreased after the additions of P and N.
3. Compared with heterotrophic bacterioplankton, picophytoplankton were most successful at the clearwater end of the lake water colour gradient. Phytoplankton dominated over heterotrophic bacteria in the clearwater systems possibly because heterotrophic bacteria in such lakes are dependent on organic carbon produced by phytoplankton.
4. Compared with other phytoplankton, picophytoplankton did best at intermediate DOC concentrations; flagellates dominated in the humic lakes and large autotrophic phytoplankton in the clearwater lakes.
5. Picophytoplankton were not better competitors than large phytoplankton in situations when heterotrophic bacteria had access to a non-algal carbon source. Neither did their small size lead to picophytoplankton dominance over large phytoplankton in the clearwater lakes. Possible reasons include the ability of larger phytoplankton to float or swim to reduce sedimentation losses and to acquire nutrients by phagotrophy.     

The North Atlantic Oscillation and plankton dynamics in two European lakes –- two variations on a general theme

Long‐term data on water temperature, phytoplankton biovolume, Bosmina and Daphnia abundance and the timing of the clear‐water phase were compared and analysed with respect to the influence of the North Atlantic Oscillation (NAO) in two strongly contrasting lakes in central Europe. In small, shallow, hypertrophic Müggelsee, spring water temperatures and Daphnia abundance both increased more rapidly than in large, deep, meso/oligotrophic Lake Constance. Because of this, the clear‐water phase commenced approximately three weeks earlier in Müggelsee than in Lake Constance. In Müggelsee, the phytoplankton biovolume during late winter/early spring was related to the NAO index. In Lake Constance, where phytoplankton growth was inhibited by intense downward mixing during all years studied, this was not the case. However, in both lakes, interannual variability in water temperature, in Daphnia spring population dynamics and in the timing of the clear‐water phase, were all related to the interannual variability of the NAO index. The Daphnia spring population dynamics and the timing of the clear‐water phase appear to be synchronized by the NAO despite large differences between the lakes in morphometry, trophic status and flushing and mixis regimes, and despite the great distance between the lakes (～700 km). This suggests that a great variety of lakes in central Europe may possibly have exhibited similar interannual variability during the last 20 years.     

Limnology and trophic status of glacial lakes in the tropical Andes (Cajas National Park,Ecuador)

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Water temperature and stratification depth independently shift cardinal events during plankton spring succession

In deep temperate lakes, the beginning of the growing season is triggered by thermal stratification, which alleviates light limitation of planktonic producers in the surface layer and prevents heat loss to deeper strata. The sequence of subsequent phenological events (phytoplankton spring bloom, grazer peak, clearwater phase) results in part from coupled phytoplankton–grazer interactions. Disentangling the separate, direct effects of correlated climatic drivers (stratification‐dependent underwater light climate vs. water temperature) from their indirect effects mediated through trophic feedbacks is impossible using observational field data, which challenges our understanding of global warming effects on seasonal plankton dynamics. We therefore manipulated water temperature and stratification depth independently in experimental field mesocosms containing ambient microplankton and inocula of the resident grazer Daphnia hyalina. Higher light availability in shallower surface layers accelerated primary production, warming accelerated consumption and growth of Daphnia, and both factors speeded up successional dynamics driven by trophic feedbacks. Specifically, phytoplankton peaked and decreased earlier and Daphnia populations increased and peaked earlier at both shallower stratification and higher temperature. The timing of ciliate dynamics was unrelated to both factors. Volumetric peak densities of phytoplankton, ciliates and Daphnia in the surface layer were also unaffected by temperature but declined with stratification depth in parallel with light availability. The latter relationship vanished, however, when population sizes were integrated over the entire water column. Overall our results suggest that, integrated over the entire water column of a deep lake, surface warming and shallower stratification independently speed up spring successional events, whereas the magnitudes of phytoplankton and zooplankton spring peaks are less sensitive to these factors. Therefore, accelerated dynamics under warming need not lead to a trophic mismatch (given similar grazer inocula at the time of stratification). We emphasize that entire water column dynamics must be studied to estimate global warming effects on lake ecosystems.     

Phytoplankton–zooplankton seasonal timing and the 'clear-water phase' in some English lakes

SUMMARY 1. An examination is made of the relative seasonal timing of the postwinter increase of phytoplankton and zooplankton populations in four English lake basins. It centres upon weekly sampling over 20 years and rough counts of larger Crustacea, as copepods and cladocerans, from filtered samples that were used for chlorophyll a (Chl) estimation. 2. Typically, a spring maximum of phytoplankton, dominated by diatoms and earlier in the shallower lakes, is accompanied or followed by a maximum of copepods and then one of cladocerans dominated by the Daphnia hyalina–galeata complex. Regarding timing, the maximum of copepods has no apparent relation with phytoplankton abundance (Chl). The maximum of cladocerans appears to be largely independent of variation in the phytoplankton maximum, but is generally associated with a minimum in Chl. Evidence for some direct causality in this inverse correlation after the spring phytoplankton maximum is best displayed by the shallow Esthwaite Water in which the peaks of Chl and cladocerans are separated further than in the deep Windermere basins where phytoplankton growth is delayed. In Esthwaite Water, and possibly often in Windermere, a principal minimum in Chl is ascribable to grazing by Daphnia. 3. The typical inverse relationship of Chl and cladocerans is lost in some years when relatively inedible large phytoplankters (e.g. colonial chrysomonads, filamentous cyanophytes) are abundant and Chl minima are less pronounced, although maxima of cladocerans still occur. Conversely, available edible phytoplankters include various small forms grouped as μ‐algae and Cryptomonas spp.; their probable depletions by Daphnia appear to be sequential and may limit the latter's maxima, whose inception is temperature‐dependent. 4. The spring–summer maxima of cladocerans and minima of Chl are generally coincident with a main seasonal maximum of Secchi disc transparency and light penetration – to which removal of non‐phytoplankton particles by filtering cladocerans may contribute.     

Interannual variability in phytoplankton biomass and species composition in a subtropical reservoir

• 1 The interannual variability of the dominant phytoplankton populations is described in a subtropical reservoir in Queensland using weekly data for a 16-year period between 1978 and 1994. North Pine Dam, Brisbane, is in an area characterized by strong interannual variability in rainfall. This variability is linked to El Nino Southern Oscillation (ENSO) events. Between 1978 and 1994 periods of drought (during strong ENSO events) were interspersed by periods of flooding rains. Rainfall on the catchment and temperature and oxygen within the dam showed strong 40-day periodicities which also varied in strength interannually in response to ENSO events. Similar patterns of fluctuations in the 40-day periodicity have been found elsewhere in SE Australia. Seasonal cycles of stratification in the dam were a function of both hydrographic and hydrological events. Intermittent rain storms caused partial turnovers and large outflows. As much as 90% of the dam volume was exchanged in a single flood event.
• 2 The dominant phytoplankton species were similar to those frequently found in tropical and subtropical lakes and reservoirs. The phytoplankton community switched between cyanobacterial blooms (Cylindrospermopsis, Microcystis) during drought and falling water levels and diatom blooms (Aulacoseira) in response to inflows and seasonal turnovers. There appeared to be a subtle interaction between inflows, water column stability, the periodic overturns and the occurrence of the dominant species. All the dominant species showed long periods (2–4 years) of exponential increase or decrease superimposed on top of the seasonal fluctuations in abundance. These patterns of abundance led to marked interannual variability in the phytoplankton biomass. Climate variability had a major impact on the seasonal and interannual changes of the dominant phytoplankton species.
• 3 Phytoplankton biomass tended to be depressed for about 3 months after individual storm events but the data also displayed long-term lag effects (2–4 years) which destroyed any significant correlation between water residence time and biomass. Summer maxima of biomass dominated by cyanobacteria disappeared between 1985 and 1990 and were replaced by smaller winter peaks. The data presented here are not capable of unequivocally identifying the precise reason for these longer-term effects. Because of the implications for water quality management in subtropical and tropical reservoirs they warrant further study.

     

Diet shifts and food selection of perch Perca fluviatilis and roach Rutilus rutilus in humic lakes of varying water colour

The field data from four humic lakes suggested that water colour may have both direct and indirect effects on inter‐ and intra‐specific interactions of perch Perca fluviatilis and roach Rutilus rutilus. The results agree with suggestions that, compared with R. rutilus, P. fluviatilis may be an inferior forager on zooplankton in highly coloured water. As an indirect effect, water colour decreases the coverage of macrophytes and limits suitable littoral habitats, benefiting R. rutilus over P. fluviatilis. Perca fluviatilis benefiting from complex habitats does not have the advantage in macrophyte‐poor highly coloured water.     

Spring phytoplankton of 54 small lakes in southern Finland

The abundance and species composition of phytoplankton communities were studied rapidly following the spring ice-melt in 54 small Finnish lakes that form a unique mosaic of water bodies. Phytoplankton biomass and cell density varied among the study lakes with a factor 100 between the lowest and highest values. Highest biomass and densities of phytoplankton characterized small ( < 0.05 km²) lakes with moderate or high water colour (> 80 mg Pt l^–1). In contrast, biomass was low in clear-water lakes and lakes where water throughflow was strong. Typically one species dominated most phytoplankton communities, and usually comprised up to about 45% of the total phytoplankton biomass. Two-thirds of the 103 taxa observed were Chrysophyceans and Chlorophyceans. The most common taxa wereChlamydomonas spp. (Chlorophyceae) andCryptomonas ovata (Cryptophyceae).     

Autotrophic and Heterotrophic Picoplankton in Wetlands: Differences with Lake Patterns

This study describes the occurrence, importance and seasonal patterns of picoplankton in two wetlands (TDNP and La Safor), and compares them to a system of fifteen interconnected lakes (Ruidera). In TDNP we performed a six‐year monthly study in three sites of the wetland. Bacterial abundance increased throughout time and the autotrophic picoplankton (APP) range was wide (up to 33 × 10⁶ cells/ml). The annual averaged APP contribution to total picoplankton and phytoplankton biovolumes was 0.5–22% and 0.03–6% respectively. There were large differences among sites in terms of APP absolute and relative abundance and seasonal patterns. In La Safor, the APP relative contribution to picoplankton and phytoplankton biovolumes was 0–25% and 0–40%, respectively, while in the Ruidera lakes was 0–47% and 0–5%, respectively. In the three systems there was a significant correlation between bacterial abundance and chlorophyll a but the slopes of the linear regressions were different. No significant relationships were found of APP abundance and trophic status in the wetlands, but were noted in the lake system. There was no clear relationship of APP contribution to total phytoplankton biomass to the trophic gradient in wetlands. In the lakes, the higher contribution of APP was found in those with higher trophic levels.     

Vertical Distribution of Potentially Pathogenic Free-Living Amoebae in Freshwater Lakes1

The vertical distribution of thermotolerant (37°C and 45°C) free-living amoebae (FLA) in warm monomictic lakes was determined in relation to the onset of thermal stratification and associated physical and chemical changes. The position of abiotic or biotic paniculate layers in the water column was located by using a submersible horizontal beam transmissometer that measures attenuance, or the absorption and scattering of light by participates in the water column. During mixis, the vertical distribution of amoebae was sporadic with significant numbers of FLA only occurring in clay layers caused by runoff after heavy rains. With the onset of thermal stratification in the lakes, phytoplankton layers began to form. Few amoebae were isolated from layers containing flagellated phytoplankton; however, significant (P < 0.005) numbers of FLA were isolated from two paniculate layers dominated by the filamentous blue-green algae Aphanizomenon and Lyngbya, respectively. By late June, a persistent detrital or decomposition layer formed in the lower metalimnion, as well as a hypolimnetic iron layer where the Fe²⁺ state was predominant. In this midsummer period, 13 Naegieria fowleri were isolated, with three from the detrital layer and seven from the iron layer. The presence of attenuation zones was found to be the best indicator of the vertical distribution of FLA in the water column, and such layers represent an important, previously undescribed habitat for potentially pathogenic FLA.     

Drivers of interannual variability in virioplankton abundance at the coastal western Antarctic peninsula and the potential effects of climate change

An 8‐year time‐series in the Western Antarctic Peninsula (WAP) with an approximately weekly sampling frequency was used to elucidate changes in virioplankton abundance and their drivers in this climatically sensitive region. Virioplankton abundances at the coastal WAP show a pronounced seasonal cycle with interannual variability in the timing and magnitude of the summer maxima. Bacterioplankton abundance is the most influential driving factor of the virioplankton, and exhibit closely coupled dynamics. Sea ice cover and duration predetermine levels of phytoplankton stock and thus, influence virioplankton by dictating the substrates available to the bacterioplankton. However, variations in the composition of the phytoplankton community and particularly the prominence of Diatoms inferred from silicate drawdown, drive interannual differences in the magnitude of the virioplankton bloom; likely again mediated through changes in the bacterioplankton. Their findings suggest that future warming within the WAP will cause changes in sea ice that will influence viruses and their microbial hosts through changes in the timing, magnitude and composition of the phytoplankton bloom. Thus, the flow of matter and energy through the viral shunt may be decreased with consequences for the Antarctic food web and element cycling.     

Essential fatty acids and phosphorus in seston from lakes with contrasting terrestrial dissolved organic carbon content

1. It is often assumed that lakes highly influenced by terrestrial organic matter (TOM) have low zooplankton food quality because of elemental and/or biochemical deficiencies of the major particulate organic carbon pools. We used the biochemical [polyunsaturated fatty acids, especially eicosapentaenoic acid (EPA) – 20:5ω3] and elemental (C : P ratio) composition of particulate matter (PM) as qualitative measures of potential zooplankton food in two categories of lakes of similar primary productivity, but with contrasting TOM influence (clear water versus humic lakes). 2. C : P ratios (atomic ratio) in PM were similar between lake categories and were above 400. The concentration (μg L⁻¹) and relative content (μg mg C⁻¹) of EPA, as well as the particulate organic carbon concentration, were higher in the humic lakes than in the clear‐water lakes. 3. Our results show high fatty acid quality of PM in the humic lakes. The differences in the biochemical quality of the potential zooplankton food between lake categories can be attributed to the differences in their phytoplankton communities. 4. High biochemical quality of the food can result in high efficiency of energy transfer in the food chain and stimulate production at higher trophic levels, assuming that zooplankton are able to ingest and digest the resource available.     

Seasonal variation in phytoplankton composition and physical-chemical features of the shallow Lake Doïrani,Macedonia, Greece

Phytoplankton species composition, seasonal dynamics and spatial distribution in the shallow Lake Doïrani were studied during the growth season of 1996 along with key physical and chemical variables of the water. Weak thermal stratification developed in the lake during the warm period of 1996. The low N:P ratio suggests that nitrogen was the potential limiting nutrient of phytoplankton in the lake. In the phytoplankton of the lake, Chlorophyceae were the most species-rich group followed by Cyanophyceae. The monthly fluctuations of the total phytoplankton biomass presented high levels of summer algal biomass resembling that of other eutrophic lakes. Dinophyceae was the group most represented in the phytoplankton followed by Cyanophyceae. Diatomophyceae dominated in spring and autumn. Nanoplankton comprised around 90% of the total biomass in early spring and less than 10% in summer. The seasonal dynamics of phytoplankton generally followed the typical pattern outlined for other eutrophic lakes. R-species (small diatoms), dominant in the early phase of succession, were replaced by S-species (Microcystis, Anabaena, Ceratium) in summer. With cooling of the water in September, the biomass of diatoms (R-species) increased. The summer algal maxima consisted of a combination of H and M species associations (sensu Reynolds). Phytoplankton development in 1996 was subject to the combined effect of the thermal regime, the small depth of mixing and the increased sediment-water interactions in the lake, which caused changes in the underwater light conditions and nutrient concentrations.     

Variable littoral‐pelagic coupling as a food‐web response to seasonal changes in pelagic primary production

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Palaeoenvironmental significance of the green alga Botryococcus in the lacustrine rotliegend (upper carboniferous ‐ lower permian)

The green alga Botryococcus is an important component of the phytoplankton in lake sediments from the Rotliegend of Central Europe. The first microscopic fossils from the Saar‐Nahe Basin that can be assigned to this genus are described and illustrated here. Their distribution at various stratigraphic levels shows that the primary production of the lakes was at least partially controlled by these green algae. The frequency of fossils in the sediments is correlated with grain size, which varied with the depositional environment. Their abundance is greater in sediments deposited in shallow water, especially in small lakes and in the littoral environments of larger lakes. Sediments of deep‐water lake bottoms contain only minor amounts of algal remains. The abundance of kerogen originating from phytoplankton and occurrences of Botryococcus fossils are correlated in these lake deposits.     

Water colour, phosphorus and alkalinity are the major determinants of the dominant phytoplankton species in European lakes

Analysis of phytoplankton data from about 1,500 lakes in 20 European countries has revealed that two-thirds of the species that dominate lakes during the summer are dominant right across Europe. Using Canonical Correspondence Analyses, we have examined how both habitat conditions within lakes and environmental factors over broad geographical scales explained the distribution of the 151 most common summer dominant species. The distributions of these species were best explained by water colour and latitude, although alkalinity and total phosphorus also appeared to be important explanatory factors. Contrary to our original hypothesis, summer water temperatures had a negligible impact on the distribution of dominants, although, due to the restricted summer season we examined, only a limited temperature gradient was present in the dataset. Cryptophytes occurred more frequently among dominants in Northern Europe whereas cyanobacteria and dinophytes dominated more in Central and Southern Europe. Our analyses suggest that besides nutrient concentrations, other water chemistry variables, such as alkalinity and the content of humic substances, have at least as important a role in determining the distribution of the dominant phytoplankton species in European lakes.     

Deep chlorophyll maxima and UVR acclimation by epilimnetic phytoplankton

1. It is well established that ultraviolet radiation (UVR) has many harmful effects on phytoplankton, but the factors controlling algal sensitivity to UVR are not fully understood. 2. We exposed phytoplankton communities from the epilimnia and deep chlorophyll maxima (DCM) of 2 Canadian lakes to 14 irradiance treatments of various spectral quality and monitored changes in the maximum quantum efficiency of Photosystem II photochemistry (F_v/F_m) using a pulse amplitude modulation fluorometer. 3. Phytoplankton from DCM did not show marked differences from epilimnetic communities in taxonomy or nutrient status, but exhibited substantially higher photosynthetic impairment under UVR exposure. 4. Our results suggest that epilimnetic phytoplankton acclimate to in situ light conditions in a spectrally‐specific manner, and that ultraviolet‐A radiation is a stronger stressor than ultraviolet‐B or photosynthetically active radiation in the mixed layers of our study lakes. Model estimates of damage and recovery rate constants revealed that the phytoplankton of the two lakes relied upon different strategies of UVR‐acclimation, in one lake minimising susceptibility to photodamage and in the other maximising recovery efficiency.     

Filamentous green algae inhibit phytoplankton with enhanced effects when lakes get warmer

1. Filamentous green algae (FGA) may represent an alternative state in high‐nutrient shallow temperate lakes. Furthermore, a clear water state is sometimes associated with the dominance of FGA; however, the mechanisms involved remain uncertain. 2. We hypothesised that FGA may promote a clear water state by directly suppressing phytoplankton growth, mostly via the release of allelochemicals, and that this interaction may be affected by temperature. 3. We examined the relationships between FGA, phytoplanktonic chlorophyll a concentrations and zooplankton in a series of mesocosms (2.8 m³) mimicking enriched shallow ponds now and in a future warmer climate (0 and c. 5 °C above ambient temperatures). We then tested the potential allelopathic effects of FGA (Cladophora sp. and Spirogyra sp.) on phytoplankton using several short‐term microcosms and laboratory experiments. 4. Mesocosms with FGA evidenced lower phytoplanktonic chlorophyll a concentrations than those without. Zooplankton and zooplankton : phytoplankton biomass ratios did not differ between mesocosms with and without FGA, suggesting that grazing was not responsible for the negative effects on phytoplanktonic biomass (chlorophyll a). 5. Our field microcosm experiments demonstrated that FGA strongly suppressed the growth of natural phytoplankton at non‐limiting nutrient conditions and regardless of phytoplankton initial concentrations or micronutrients addition. Furthermore, we found that the negative effect of FGA on phytoplankton growth increased up to 49% under high incubation temperatures. The experiment performed using FGA filtrates confirmed that the inhibitory effect of FGA on phytoplankton may be attributed to allelochemicals. 6. Our results suggest that FGA control of phytoplankton growth may be an important mechanism for stabilising clear water in shallow temperate lakes dominated by FGA and that FGA may play a larger role when lakes get warmer.