Nutrient ratios and phytoplankton community structure in the large, shallow, eutrophic, subtropical Lakes Okeechobee (Florida, USA) and Taihu (China)

Analysis of ten- and four-year datasets for the large, shallow, subtropical, and eutrophic Lakes Okeechobee (USA) and Taihu (China), respectively, suggest that resource-ratio explanations for cyanobacteria dominance may not apply to these two lakes. Datasets were examined to identify relationships between nutrient ratios [total nitrogen (TN):total phosphorus (TP) and ammonium (NH₄ ⁺):oxidized N (NO _x )] and phytoplankton community structure (as proportions of cyanobacteria and diatoms to total phytoplankton biomass). Datasets were pooled by sampling month, averaged lake-wide, and analyzed with linear regression. In Okeechobee, the cyanobacteria proportion increased and the diatom proportion decreased with increasing TN:TP. In Taihu, cyanobacteria decreased with increasing TN:TP, but the opposite trend observed for diatoms was marginally significant. Okeechobee cyanobacteria increased and diatoms decreased with increasing NH₄ ⁺:NO _x , but no significant relationships between phytoplankton and NH₄ ⁺:NO _x were observed in Taihu. Both lakes had significant relationships between phytoplankton community structure and total nutrients, but these relationships were the opposite of those expected. Relationships between phytoplankton community structure and water quality parameters from the previous month resulted in improved relationships, suggesting a predictive capability. Statistical analysis of the entire datasets (not pooled) supported these and additional relationships with other parameters, including temperature and water clarity.     

Increased risk of cyanobacterial blooms in northern high‐latitude lakes through climate warming and phosphorus enrichment

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Acceleration of cyanobacterial dominance in north temperate‐subarctic lakes during the Anthropocene

Increases in atmospheric temperature and nutrients from land are thought to be promoting the expansion of harmful cyanobacteria in lakes worldwide, yet to date there has been no quantitative synthesis of long‐term trends. To test whether cyanobacteria have increased in abundance over the past ~ 200 years and evaluate the relative influence of potential causal mechanisms, we synthesised 108 highly resolved sedimentary time series and 18 decadal‐scale monitoring records from north temperate‐subarctic lakes. We demonstrate that: (1) cyanobacteria have increased significantly since c. 1800 ce , (2) they have increased disproportionately relative to other phytoplankton, and (3) cyanobacteria increased more rapidly post c. 1945 ce . Variation among lakes in the rates of increase was explained best by nutrient concentration (phosphorus and nitrogen), and temperature was of secondary importance. Although cyanobacterial biomass has declined in some managed lakes with reduced nutrient influx, the larger spatio‐temporal scale of sedimentary records show continued increases in cyanobacteria throughout the north temperate‐subarctic regions.     

Long-term lake acidification trends in high- and low-sulphate deposition regions from Nova Scotia,Canada

Diatom-based paleolimnological techniques were used to study 14 lakes from two regions of Nova Scotia which represent regions of high and low sulphate deposition. Using decadal scale intervals, changes in diatom assemblages and diatom-inferred pH were tracked in relation to deposition of anthropogenic-sourced strong acids. Eight study lakes were located in Kejimkujik National Park in the southwestern part of the province, which receives an annual sulphate deposition (2000–2002) of ~10.5 kg ha⁻¹ yr⁻¹. These lakes showed significant changes in diatom assemblages with overall diatom-inferred acidification of ~0.5 pH units starting between 1925 and 1940, with the timing of acidification related to pre-industrial (or pre−1850) lakewater pH. Six study lakes were located in Cape Breton Highlands National Park, in northern Nova Scotia, a region of lower sulphate deposition. These lakes did not show any consistent trends in diatom assemblages or inferred pH values consistent with recent acidic deposition, but rather variations that may be related to climatic influences. Nova Scotia lakes that have been most impacted by acidic deposition had the lowest pre-industrial lakewater pH values and were in an area of relatively high sulphate deposition. Handling editor: K. Martens     

Nitrogen deposition and warming – effects on phytoplankton nutrient limitation in subarctic lakes

The aim of this study was to predict the combined effects of enhanced nitrogen (N) deposition and warming on phytoplankton development in high latitude and mountain lakes. Consequently, we assessed, in a series of enclosure experiments, how lake water nutrient stoichiometry and phytoplankton nutrient limitation varied over the growing season in 11 lakes situated along an altitudinal/climate gradient with low N‐deposition (<1 kg N ha^?1 yr^?1) in northern subarctic Sweden. Short‐term bioassay experiments with N‐ and P‐additions revealed that phytoplankton in high‐alpine lakes were more prone to P‐limitation, and with decreasing altitude became increasingly N‐ and NP‐colimited. Nutrient limitation was additionally most obvious in midsummer. There was also a strong positive correlation between phytoplankton growth and water temperature in the bioassays. Although excess nutrients were available in spring and autumn, on these occasions growth was likely constrained by low water temperatures. These results imply that enhanced N‐deposition over the Swedish mountain areas will, with the exception of high‐alpine lakes, enhance biomass and drive phytoplankton from N‐ to P‐limitation. However, if not accompanied by warming, N‐input from deposition will stimulate limited phytoplankton growth due to low water temperatures during large parts of the growing season. Direct effects of warming, allowing increased metabolic rates and an extension of the growing season, seem equally crucial to synergistically enhance phytoplankton development in these lakes.     

Identification and quantification of phytoplankton groups in lakes using new pigment ratios – a comparison between pigment analysis by HPLC and microscopy

1. Pigment analysis by high‐performance liquid chromatography (HPLC) combined with data analysis using the CHEMTAX program has proven to be a fast and precise method for determining the abundance of phytoplankton groups in marine environments. To determine whether CHEMTAX is applicable also to freshwater phytoplankton, 20 different species of freshwater algae were cultured and their pigment/chlorophyll a (Chl a) ratios determined for exponential growth at three different light intensities and for stationary growth at one light intensity. 2. The different treatments had a relatively insignificant impact on the absolute values of the diagnostic pigment/Chl a ratios, with the exception of cyanobacteria and cryptophytes for which the zeaxanthin/Chl a and alloxanthin/Chl a ratios varied considerably. 3. The pigment ratios were tested on samples collected in six different eutrophic Danish lakes during two summer periods using the CHEMTAX program to calculate the biomass of the phytoplankton groups as Chl a. The CHEMTAX‐derived seasonal changes in Chl a biomass corresponded well with the volume of the microscopically determined phytoplankton groups. More phytoplankton groups were detected by the pigment method than by the microscopic method. 4. Applying the pigment ratios developed in this study, the pigment method can be used to determine the abundance of the individual phytoplankton groups, which are useful as biological water quality indicators when determining the ecological status of freshwater lakes.     

Hydrology and Diatom Phytoplankton of High Arctic Lakes and Ponds on Store Koldewey,Northeast Greenland

We document hydrological and phytoplankton characteristics of nine lakes and two ponds on Store Koldewey, a culturally undisturbed island off Northeast Greenland. The limnological survey included the recording of temperature, conductivity, oxygen concentration and saturation, pH, ionic composition, transparency, and the diatom phytoplankton community. In summer 2003, the lakes were cold, monomictic, thermally unstratified, alkaline and likely oligotrophic water bodies. Diatom phytoplankton was present in six lakes and consisted of four dominant species (Aulacoseira tethera, Cyclotella pseudostelligera, C. rossii, and Fragilaria tenera). The concentration of planktonic diatoms varied distinctly between the lakes. (© 2005 WILEY‐VCH Verlag GmbH & Co. KGaA, Weinheim)     

ALLELOPATHIC GROWTH INHIBITION OF SELECTED PHYTOPLANKTON SPECIES BY SUBMERGED MACROPHYTES1

Allelopathic effects of submerged macrophytes on the growth and photosynthesis of different unialgal cultures of planktonic cyanobacteria, a diatom, and a green alga were tested in coexistence experiments using dialysis cultures. The method applied allowed measurements under conditions similar to that in lakes but without nutrient and light limitation. Growth and photosynthesis were measured with a pulse amplitude modulated fluorometer as an increase of chl a fluorescence and activity of PSII, respectively. Eurasian water milfoil Myriophyllum spicatum L. and rigid hornwort Ceratophyllum demersum L. proved to inhibit the PSII activity and then growth of the investigated phytoplankton species, whereas sago pondweed Potamogeton pectinatus L. showed no effect. Growth inhibition was dependent on biomass of M. spicatum. Considerable differences between phytoplankton groups and among species of cyanobacteria were found regarding their response to M. spicatum. Members of the Oscillatoriales and Microcystis aeruginosa Kütz. emend. Elenkin were more sensitive than the cyanobacterium Aphanizomenon flos‐aquae Ralfs ex Born. et Flah., the diatom Stephanodiscus minutulus (Kütz) Cleve et Möller, and the green alga Scenedesmus armatus Chodat. A possible contribution of this result to changes in the phytoplankton succession of lakes after loss of macrophytes is discussed.     

Freshwater diatom biogeography in the Canadian Arctic Archipelago

Aim Document the biogeographical distributions of diatoms in the Canadian Arctic in relation to environmental factors. Location The Canadian Arctic Archipelago. Methods Diatoms were extracted from lake sediments and treated using standard methods. Rarefaction‐estimated species richness, diatom concentrations (valves cc^?1), and diatom abundance were mapped using a Geographic Information System. The physical and chemical parameters of the lakes were measured. Results A total of 326 taxa from 63 genera were found in 62 lakes of the study area. Up to 85 and as low as eight taxa were identified in any one lake, and rarefaction‐estimated species richness correlated with lake size. Nearby lakes could have greatly different diatom assemblages. Many taxa showed limited distributions. Response surfaces and ordination techniques indicate that a large number of taxa prefer colder regions of the Arctic while the abundance of others was influenced by lake water chemical parameters such as TKN and SiO₂. Main conclusions Although many taxa were common and found across the study area, diatom assemblages showed regional differences in the Arctic. Some taxa were not found in either the northern or southern parts of the Archipelago and others were restricted to particular regions. Newly delineated genera showed interpretable geographical patterns and could be related to environmental factors, suggesting that this more natural grouping may enhance our understanding of diatom ecology. Geographical, physical, and chemical factors are needed to explain diatom distributions in the Arctic.     

CYANOBACTERIAL DOMINANCE OF POLAR FRESHWATER ECOSYSTEMS: ARE HIGH-LATITUDE MAT-FORMERS ADAPTED TO LOW TEMPERATURE?1

Although it is generally believed that cyanobacteria have high temperature optima for growth (> 20° C), mat-foming cyanobacteria are dominant in many types of lakes, streams, and ponds in the Arctic and Antarctic. We studied the effect of temperature on growth (μ) and relative pigment composition of 27 isolates of cyanobacteria (mat-forming Oscillatoriaceae) from the Arctic, subarctic, and Antarctic to investigate whether they are a) adapted to the low temperature (i.e. psychrophilic) or b) tolerant of the low temperature of the polar regions (i.e. psychrotrophic). We also derived a parabolic function that describes both the rise and the decline of cyanobacterial growth rates with increasing temperature. The cyanobacteria were cultured at seven different temperatures (5°-35° C at 5° C intervals), with continuous illumination of 225 μmol photons.m⁻².s⁻¹. The parabolic function fits the μ-temperature data with 90% confidence for 75% of the isolates. Among the 27 isolates of cyanobacteria studied, the temperature optima (T_opt) for growth ranged from 15° to 35° C, with an average of 19.9° C. These results imply that most polar cyanobacteria are psychrotrophs, not psychrophiles. The cyanobacteria grew over a wide temperature range (typically 20° C) but growth rates were low men at T_opt (average μ_max of 0.23 ± 0.069 d⁻¹). Extremely slow growth rates at low temperature and the high temperature for optimal growth imply that the cyanobacteria are not adapted genetically to cold temperatures, which characterize their ambient environment. Other competitive advantages such as tolerance to desiccation, freeze—thaw cycles, and bright, continuous solar radiation may contribute to their dominance in polar aquatic ecosystems.     

Determining algal assemblages in oligotrophic lakes and streams: comparing information from newly developed pigment/chlorophyll a ratios with direct microscopy

1. Pigment analyses by high performance liquid chromatography (HPLC) are commonly used for determining algal groups in marine and estuarine areas but are underdeveloped in freshwaters. In this study, 15 characteristic pelagic algal species (representing five algal groups) of oligo‐ / mesotrophic lakes were cultured and pigment / Chl a ratios determined at three light intensities. 2. With the exception of cyanophytes, light treatment had little effect on pigment / Chl a ratios. This justifies the use of the same pigment / Chl a ratios during seasonal studies where light conditions may change. 3. The determined pigment / Chl a ratios were tested on seasonal samples from five oligo‐ / mesotrophic lakes and three streams using CHEMTAX software. Pigment ratios of both pelagic and benthic algal communities from the lakes and streams were analysed to determine whether the pelagic algae‐based ratios can be used for benthic algal communities. 4. HPLC combined with CHEMTAX was useful for identifying freshwater phytoplankton classes and for quantifying the abundance of phytoplankton groups. However, although correlations were significant for six of seven phytoplankton classes studied, they were weak and varied with season. 5. HPLC was valid for quantifying benthic diatom groups in stream samples, whereas for lakes more benthic algal groups were recorded with HPLC than with microscopy and correlations between the two methods were not significant. 6. The use of both HPLC and microscopy is recommended as a cost‐efficient method for analysing many samples. It is crucial, however, that the CHEMTAX software is calibrated with the correct information, and the user is aware of the limitations.     

Clustering and canonical correspondence analysis of phytoplankton and environmental variables in Swedish lakes

Phytoplankton data consisting of 145 species from a limnological study of lakes from relatively undisturbed areas throughout Sweden were analysed in relation to 11 physical and chemical environmental variables. Three multivariate methods were applied: WPGMA clustering and TWINSPAN for classification, and detrended canonical correspondence analysis (DCCA), a recent technique which extracts ordination axes that can be related directly to variation in the environment. Three types of lakes were recognized consistently: acid humic lakes with Gonyostomum semen as the dominant species, very acid impoverished lakes with rather few, stress-tolerant species, and subarctic lakes with low total biomass but with a varied phytoplankton flora. DCCA allowed a straightforward display of the locations of lakes and species along environmental gradients (including the acidification gradient) reflected in phytoplankton composition. It is suggested that such analyses may be a useful tool for the early detection of environmental change.     

Diatom assemblages and their relationships to lakewater nitrogen levels and other limnological variables from 36 lakes and ponds on Banks Island,N.W.T., Canadian Arctic

Banks Island, in the Canadian Arctic Archipelago, has been identified as an important reference site for studies of environmental change, especially as it relates to climatic warming. The island is logistically manageable (i.e. researchers can survey the entire island in one field season) and, most importantly, spans three major ecoclimatic regions supporting a diverse and large bird and mammal population. Developing upon earlier work by the authors describing the limnology of Banks Island, this current study: (1) examines which physical and chemical limnological variables influence diatom assemblages in this relatively lush island; and (2) explores variations in the diatom assemblages by ecoclimatic zones. The relationship between diatom taxa from a 36 lake/pond surface sediment calibration set and a suite of limnological variables was explored using multivariate statistics. Dominant diatom species varied based on changing limnological characteristics, particularly between the colder, ultra-oligotrophic lakes in the more northern High Arctic regions compared to the warmer, more nitrogen-rich sites in the Low Arctic regions of Banks Island. Exploration of diatom ecoclimatic and environmental preferences revealed interesting relationships, including the development of a diatom-based transfer function that could be used to track overall trends on lakewater nitrogen concentrations, which may enable future paleolimnological studies to track shifts in nutrient levels and climatic, and other environmental changes. Handling editor: J. Padisak     

Responses of phytoplankton to fish predation and nutrient loading in shallow lakes: a pan-European mesocosm experiment

1. The impacts of nutrients (phosphorus and nitrogen) and planktivorous fish on phytoplankton composition and biomass were studied in six shallow, macrophyte‐dominated lakes across Europe using mesocosm experiments. 2. Phytoplankton biomass was more influenced by nutrients than by densities of planktivorous fish. Nutrient addition resulted in increased algal biomass at all locations. In some experiments, a decrease was noted at the highest nutrient loadings, corresponding to added concentrations of 1 mg L^?1 P and 10 mg L^?1 N. 3. Chlorophyll a was a more precise parameter to quantify phytoplankton biomass than algal biovolume, with lower within‐treatment variability. 4. Higher densities of planktivorous fish shifted phytoplankton composition toward smaller algae (GALD < 50 μm). High nutrient loadings selected in favour of chlorophytes and cyanobacteria, while biovolumes of diatoms and dinophytes decreased. High temperatures also may increase the contribution of cyanobacteria to total phytoplankton biovolume in shallow lakes.     

Diatom evidence for the timing and causes of eutrophication in Lake Victoria,East Africa

The determination of the history and causes of recent eutrophication and intensified thermal stratification in Lake Victoria is still hampered by the sparsity of paleolimnologic coverage of the enormous lake. Five new diatom records from Ugandan waters now show that a transition from Aulacoseira-dominated planktonic assemblages to those dominated by long Nitzschia spp., occurred in northern coastal sites from the mid-1970s to mid-1980s. Similar transitions developed from the late 1960s to early 1970s offshore and from the 1940s to early 1950s along the Kenyan coast, suggesting a time-transgressive process. These changes are not readily attributable to the trophic effects of Nile perch population growth during the early 1980s, but more likely reflect responses to long-term nutrient enrichment and climatic instability in the region. The diversity of planktonic diatom communities has declined dramatically, and a namesake variety of Aulacoseira nyassensis may now be nearly extirpated. Although local phytoplankton communities varied considerably in the past, the current domination of diatom assemblages by Nitzschia is apparently unprecedented in the 15,000-year history of Lake Victoria.     

Warmer climates boost cyanobacterial dominance in shallow lakes

Dominance by cyanobacteria hampers human use of lakes and reservoirs worldwide. Previous studies indicate that excessive nutrient loading and warmer conditions promote dominance by cyanobacteria, but evidence from global scale field data has so far been scarce. Our analysis, based on a study of 143 lakes along a latitudinal transect ranging from subarctic Europe to southern South America, shows that although warmer climates do not result in higher overall phytoplankton biomass, the percentage of the total phytoplankton biovolume attributable to cyanobacteria increases steeply with temperature. Our results also reveal that the percent cyanobacteria is greater in lakes with high rates of light absorption. This points to a positive feedback because restriction of light availability is often a consequence of high phytoplankton biovolume, which in turn may be driven by nutrient loading. Our results indicate a synergistic effect of nutrients and climate. The implications are that in a future warmer climate, nutrient concentrations may have to be reduced substantially from present values in many lakes if cyanobacterial dominance is to be controlled.     

Climate drivers of diatom distribution in shallow subarctic lakes

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Functional groups of phytoplankton shaping diversity of shallow lake ecosystems

Phytoplankton of eutrophic shallow lakes are frequently dominated by one species or species of the same functional group, resulting in species-pure algal assemblages. Knowledge of the structure of these assemblages is essential to understand their functioning; therefore, species and functional diversity were investigated in five sub-types of eutrophic shallow lake. Among the sub-types, astatic saline lakes and hypertrophic ponds had type-specific assemblages dominated by S_N and W0, W1 codons. The diversity of the phytoplankton in the sub-types was quite similar, except for the astatic saline lakes, which were characterised by lower values of both functional and species diversity. We found that both functional and species diversity were low when bloom-forming cyanobacteria (H1, S_N functional groups) became dominant. Dominance of other groups (J, Y, L_O and W1) did not coincide with decrease in species diversity. Analysis of the biovolume versus diversity relationships revealed that decrease in diversity might be expected at biovolume >20 mm³ l⁻¹ for shallow lakes.     

Selenium as a growth factor for plankton algae in laboratory experiments and in some Swedish lakes

The growth requirements of inorganic and organic selenium (Se) for the dinoflagellate Peridinium cinctum fa. westii and the diatom Stephanodiscus hantzschii var. pusillus and eight species of green algae are demonstrated. A new mineral culture medium for P. cinctum is presented. Using P. cinctum as a test alga, bioassays were carried out on waters from a calciferous lake and from some acidified, fertilized, lime-enriched or humic lakes for determination of their contents of bioavailable forms of Se (bioactive Se). Some of the results were related to Tot-Se and selenite-Se, measured chemically, to pH and the occurrence of ambient phytoplankton. In calcium-rich Lake Erken (pH ～ 8) blooms and decline of algae coincided with decreases (from >70 to ～20 ng Se · l^?1) and increases (up to >80 ng · l^?1) of bioactive Se, indicating uptake and release of inorganic or organic forms of Se. Acidified lakes (pH < 5) generally demonstrated much lower concentrations of bioactive Se (<20 ng · l^?1) than neutral, fertilized, lime-enriched, slightly acidic or humic lakes (>20 ng · l^?1). The correlation between pH and bioactive Se was positive (r ～ 0.92; n = 7). The ecological importance of Se is suggested for some common species of phytoplankton.     

Detection of organic pollution of streams in southern Sweden using benthic macroinvertebrates

A ²¹⁰Pb-dated sediment core from a small bay in the southern basin of Lake Petén Itzá, Guatemala documents recent cultural eutrophication. Increased sediment accumulation beginning ～1930 A.D. coincided with catchment population growth and was a consequence of watershed deforestation and increased surface run-off. At the same time, geochemical records from the Lake Petén Itzá sediment core indicate increased phosphorus loading and organic matter accumulation. High nutrient concentrations after 1965 A.D. coincided with lower sediment C/N ratios, suggesting an increase in the relative contribution of phytoplankton to the organic matter pool. This inference is confirmed by the dominance of eutrophic and hypereutrophic diatom species. Organic matter δ¹³C values decreased after 1965 A.D., seemingly contradicting other indicators of recent eutrophication in the southern basin of Lake Petén Itzá. Relatively depleted δ¹³C values in recent sediments, however, may reflect a contribution from ¹³C-depleted sewage effluent. Increased δ¹⁵N of organic matter after 1965 A.D. indicates changes in the dissolved inorganic nitrogen delivered to the lake. The relatively small increase in δ¹⁵N (～0.6‰ ) is less than might be expected with nitrate loading from sewage and soils, and might be offset by the presence of nitrogen-fixing cyanobacteria with low δ¹⁵N values.