Long-term Phytoplankton Research Articles

Analysis of Long-Term Phytoplankton Dynamics with Environmental Factors in Harima-Nada, Eastern Seto Inland Sea, Japan

Analysis of Long-Term Phytoplankton Dynamics with Environmental Factors in Harima-Nada, Eastern Seto Inland Sea, Japan

Long-term dynamics and drivers of phytoplankton biomass in eutrophic Lake Taihu

Understanding the relative effect sizes of climate-related environmental variables and nutrients on the high annual variation in the phytoplankton biomass in eutrophic lakes is important for lake management efforts. In this study, we used a data set of phytoplankton dynamics in eutrophic Lake Taihu that cover more than two decades (1993–2015) to show the variation in and the drivers of phytoplankton biomass under complex, fluctuating environmental conditions. Our results showed that the phytoplankton biomass increased slowly over the studied period despite the recent decrease in nutrient levels. The distribution of the phytoplankton biomass expanded spatially towards the central lake region, and seasonally towards the autumn and winter. Nutrients were still the primary predictors of the long-term phytoplankton biomass trend. The effect size of climate-related variables was also high and close to the effect size of nutrients. Among the climate-related variables, wind speed and underwater available light were more important predictors than temperature. The biomass of the phytoplankton taxonomic groups showed different responses to the environmental variables based on their niches. However, the compensatory dynamics affecting biomass were weak at phylum level, and synchronous dynamics drove the variation in total biomass. Our findings highlight the effect of climate-related variables on the phytoplankton biomass in Lake Taihu, which has experienced high nutrient loadings and concentrations for more than two decades. Therefore, changes in climate-related variables, such as wind speed and underwater available light, should be considered when evaluating the amount that nutrients should be reduced in Lake Taihu for future lake management.

Morphology and toxicity of Pseudo-nitzschia species in the northern Benguela Upwelling System

The Benguela upwelling system, considered the world’s most productive marine ecosystem, has a long record of potentially toxic diatoms belonging to the genus Pseudo-nitzschia. Species of Pseudo-nitzschia were reported as early as 1936 from the northern Benguela upwelling system (nBUS). For the current study, long-term phytoplankton monitoring data (2004–2011) for the Namibian coast were analysed to examine inshore and offshore temporal distribution of Pseudo-nitzschia species, their diversity and ultrastructure. The potentially toxigenic P. pungens and P. australis were the dominant inshore species, whereas offshore Pseudo-nitzschia showed a higher diversity that also included potentially toxic species. During a warming event, a community shift from P. pungens and P. australis dominance to P. fraudulenta and P. multiseries was documented in the central nBUS.A case study of a toxic event (August 2004) revealed that P. australis and P. pungens were present at multiple inshore and offshore stations, coincident with fish (pilchard) and bird mortalities reported from the central part of Namibia. Toxin analyses (LC–MS/MS) of samples collected from June to August 2004 revealed the presence of particulate domoic acid (DA) in seawater at multiple stations (maximum ∼180 ng DA/L) in the >0.45 μm size-fraction, as well as detectable DA (0.12 μg DA/g) in the gut of one of two pilchard samples tested. These findings indicate that DA may have been associated with the fish and bird mortalities reported from this event in the nBUS. However, the co-occurrence of very high biomass phytoplankton blooms suggests that other explanations may be possible.

Phytoplankton dynamics in relation to seasonal variability and upwelling and relaxation patterns at the mouth of Ria de Aveiro (West Iberian Margin) over a four-year period.

From June 2004 to December 2007, samples were weekly collected at a fixed station located at the mouth of Ria de Aveiro (West Iberian Margin). We examined the seasonal and inter-annual fluctuations in composition and community structure of the phytoplankton in relation to the main environmental drivers and assessed the influence of the oceanographic regime, namely changes in frequency and intensity of upwelling events, over the dynamics of the phytoplankton assemblage. The samples were consistently handled and a final subset of 136 OTUs (taxa with relative abundance > 0.01%) was subsequently submitted to various multivariate analyses. The phytoplankton assemblage showed significant changes at all temporal scales but with an overriding importance of seasonality over longer- (inter-annual) or shorter-term fluctuations (upwelling-related). Sea-surface temperature, salinity and maximum upwelling index were retrieved as the main driver of seasonal change. Seasonal signal was most evident in the fluctuations of chlorophyll a concentration and in the high turnover from the winter to spring phytoplankton assemblage. The seasonal cycle of production and succession was disturbed by upwelling events known to disrupt thermal stratification and induce changes in the phytoplankton assemblage. Our results indicate that both the frequency and intensity of physical forcing were important drivers of such variability, but the outcome in terms of species composition was highly dependent on the available local pool of species and the timing of those events in relation to the seasonal cycle. We conclude that duration, frequency and intensity of upwelling events, which vary seasonally and inter-annually, are paramount for maintaining long-term phytoplankton diversity likely by allowing unstable coexistence and incorporating species turnover at different scales. Our results contribute to the understanding of the complex mechanisms of coastal phytoplankton dynamics in relation to changing physical forcing which is fundamental to improve predictability of future prospects under climate change.

Ecological niche partitioning of the invasive dinoflagellate Prorocentrum minimum and its native congeners in the Baltic Sea

This study analyses three decades of the peculiar bloom-formation history of the potentially toxic invasive planktonic dinoflagellates Prorocentrum minimum (Pavillard) Schiller in the SW Baltic Sea. We tested a research hypothesis that the unexpectedly long delay (nearly two decades) in population development of P. minimum prior to its first bloom was caused by competition with one or several closely related native dinoflagellate species due to ecological niche partitioning which hampered the spread and bloom-forming potential of the invader. We applied the ecological niche concept to a large, long-term phytoplankton database and analysed the invasion history and population dynamics of P. minimum in the SW Baltic Sea coastal waters using the data on phytoplankton composition, abundance and biomass. The ecological niche dimensions of P. minimum and its congener P. balticum were identified as the optimum environmental conditions for the species during the bloom events based on water temperature, salinity, pH, concentration of nutrients (PO43−; total phosphorus, TP; total nitrogen, TN; SiO44−), TN/TP-ratio and habitat type. The data on spatial distribution and ecological niche dimensions of P. minimum have contributed to the development of the “protistan species maximum concept”. High microplankton diversity at critical salinities in the Baltic Sea may be considered as a possible reason for the significant niche overlap and strong competitive interactions among congeners leading to prolonged delay in population growth of P. minimum preceding its first bloom in the highly variable brackishwater environment.

Long-Term Phytoplankton Dynamics in the Denishi Reservoir (Ukraine)

Long-Term Phytoplankton Dynamics in the Denishi Reservoir (Ukraine)

Variations of summer phytoplankton community related to environmental factors in a macro-tidal estuarine embayment, Hangzhou Bay, China

To explore the distribution and composition of phytoplankton community and their responses to environmental changes, summer net-collected phytoplankton and physicochemical parameters in the Hangzhou Bay during 2004–2010 were investigated. A total of four phyla and 84 species were identified, including 67 diatom and 12 dinoflagellate species. The dominant species constantly consisted of the diatoms, although the dominance of dinoflagellate and cyanobacteria increased recently. Due to great spatio-temporal variations in environmental factors (salinity, suspended solids, and nutrient concentration), significant heterogeneities in community compositions among different years and subregions (inner and middle sections, and bay mouth) were found based on the analyses of multidimensional scaling and similarity. Canonical correspondence analysis showed that salinity and Si/N were the main variables associated with algal assemblage. Compared with the historical data since the 1980s, eutrophication (N, P, and N/P increased with decreasing Si/N) was exacerbated drastically. Moreover, climatic forcing and human activities resulted in a series of physical alterations, including sediment retention, temperature increase, and salinity decrease as well as reduction in water exchanges. All these changes induced obvious increases in cell density and Chl-a while decreases in species diversity and diatom-dinoflagellate ratio as well as the shifting of dominant species. Therefore, the long-term phytoplankton variations were closely related to anthropogenic and climatic perturbations in the Hangzhou Bay.

Functional approach based on morphology as a model of phytoplankton variability in a subtropical floodplain lake: a long-term study

Long-term phytoplankton dynamics in a floodplain lake, between periods of limnophase and potamophase was studied using the morphology-based functional classification (MBFG). The work was carried out to test the hypothesis that the temporal distribution of MBFGs is influenced by the hydrosedimentological regime of the Parana River, and that these differences can be registered by analyzing the dominant MBFGs in the two periods. Samples were taken in an isolated floodplain lake on the Upper Parana River floodplain, Brazil, from 2000 to 2012, and water level, water temperature, conductivity, pH, dissolved-oxygen, euphotic zone, maximum depth, and nutrients were used to explain the distribution of MBFGs. 478 taxa were identified and distributed in seven MBFGs (I–VII). MBFG V (flagellates algae) and VI (diatoms) showed 100% frequency of occurrence. MBFG II was associated exclusively with the limnophase; MBFG IV and VII were associated with limnophase periods with higher pH and dissolved-oxygen content; and MBFG I, III, V, and VI were associated with limnophase and potamophase, mainly associated with transparency, nutrients, and conductivity. The MBFG approach represented the trends of each group in terms of its occurrence and biovolume, according to the hydrosedimentological regime, providing broad-scale information on changes in the phytoplankton.

Phytoplankton functional response to spatial and temporal differences in a cold and oligotrophic lake

We explored phytoplankton communities from a functional perspective in two connected but hydro-morphologically distinct basins during the ice-free period in the cold and oligotrophic Lake Tovel, Italy. Despite the absence of dispersal barriers, we expected a clear separation of the phytoplankton communities between basins based on their physical distinctions: a shallow and a deep basin with low and high water residence time, respectively. To investigate seasonal succession and spatiality, taxa were classified according to their functional groups (FGs) and traits (FTs). Relationships between functional classifications and environmental parameters were assessed by non-metric multidimensional scaling integrated with cluster analysis. Clustering of FGs and FTs was complementary and reflected different hydrological conditions of each basin: (i) more stable conditions and higher functional variation across seasons in the deep basin and (ii) less stable conditions and lower functional variation across seasons in the shallow basin. Phytoplankton functional composition evidenced how local conditions selected for corresponding functional attributes adapted to each basin’s environment. These results, together with the presence of rare, cold-tolerant taxa such as Pseudotetraedriella kamillae and Stephanocostis chantaicus, highlighted the need for long-term phytoplankton studies, and the creation of a FG that includes cold-adapted, oligotrophic taxa.

The Continuous Plankton Recorder survey: How can long-term phytoplankton datasets contribute to the assessment of Good Environmental Status?

Phytoplankton are crucial to marine ecosystem functioning and are important indicators of environmental change. Phytoplankton data are also essential for informing management and policy, particularly in supporting the new generation of marine legislative drivers, which take a holistic ecosystem approach to management. The Marine Strategy Framework Directive (MSFD) seeks to achieve Good Environmental Status (GES) of European seas through the implementation of such a management approach. This is a regional scale directive which recognises the importance of plankton communities in marine ecosystems; plankton data at the appropriate spatial, temporal and taxonomic scales are therefore required for implementation. The Continuous Plankton Recorder (CPR) survey is a multidecadal, North Atlantic–basin scale programme which routinely records approximately 300 phytoplankton taxa. Because of these attributes, the survey plays a key role in the implementation of the MSFD and the assessment of GES in the Northeast Atlantic region. This paper addresses the role of the CPR's phytoplankton time-series in delivering GES through the development and informing of MSFD indicators, the setting of targets against a background of climate change and the provision of supporting information used to interpret change in non-plankton indicators. We also discuss CPR data in the context of other phytoplankton data types that may contribute to GES, as well as explore future possibilities for the use of new and innovative applications of CPR phytoplankton datasets in delivering GES. Efforts must be made to preserve long-term time series, such as the CPR, which supply vital ecological information used to informed evidence-based environmental policy.

Size, seasonality, or salinity: What drives the protistan species maximum in the horohalinicum?

The protistan species maximum concept for the challenging zone of critical salinity 5–8 (the horohalinicum) where the large benthic organisms experience the destructive osmotic stress has developed smoothly into the understanding that in the environment with sharp salinity fluctuations community regulations and life strategies of small unicellular planktonic organisms differ substantially from those of large multicellular bottom-dwellers. This inductive supposition has initiated efforts to define the major organismal traits and environmental factors that might be responsible for the new biodiversity pattern. There also emerged a need to better relate the novel conceptual assertions with the ‘field/pooled data paradox’ – a controversy when the pooled long-term large-scale datasets provide results which seem to contradict the field observations. This paper uses a long-term (1972–2006) phytoplankton dataset from the Baltic Sea for testing three new compatible research hypotheses by means of correlation analysis, non-metric multi-dimensional scaling using a Bray–Curtis dissimilarity matrix, and the rarefaction analysis. Hypothesis 1 argues that the smaller and therefore the faster evolving unicellular organisms are taxonomically more diverse in the horohalinicum than in the other sectors of the salinity gradient. Hypothesis 2 suggests that the recently discovered biodiversity pattern may be caused by a pronounced seasonality within the horohalinicum which might promote the shift in plankton community composition towards dominance of small-sized species in the critical salinity regions. Hypothesis 3 proposes that the protistan species maximum in the horohalinicum is largely backed up by the significant between-sample variation in highly changeable brackish-water environment rather than by within-sample species richness. Our results provided statistical proof that algal cell size minimum underpins the protistan species maximum in the horohalinicum, and supported all three research hypotheses.

Retention time generates short-term phytoplankton blooms in a shallow microtidal subtropical estuary

In this study it was hypothesised that increasing water retention time promotes phytoplankton blooms in the shallow microtidal Patos Lagoon estuary (PLE). This hypothesis was tested using salinity variation as a proxy of water retention time and chlorophyll a for phytoplankton biomass. Submersible sensors fixed at 5 m depth near the mouth of PLE continuously measured water temperature, salinity and pigments fluorescence (calibrated to chlorophyll a) between March 2010 and 12th of December 2011, with some gaps. Salinity variations were used to separate alternating patterns of outflow of lagoon water (salinity <8; 46% of the time) and inflow of marine water (salinity >24; 35% of the time). The two transition phases represented a rapid change from lagoon water outflow to marine water inflow and a more gradually declining salinity between the dominating inflow and outflow conditions. During the latter of these, a significant chlorophyll a increase relative to that expected from a linear mixing relationship was observed at intermediate salinities (10–20). The increase in chlorophyll a was positively related to the duration of the prior coastal water inflow in the PLE. Moreover, chlorophyll a increase was significantly higher during austral spring-summer than autumn-winter, probably due to higher light and nutrient availability in the former. Moreover, the retention time process operating on time scales of days influences the long-term phytoplankton variability in this ecosystem. Comparing these results with monthly data from a nearby long-term water quality monitoring station (1993–2011) support the hypothesis that chlorophyll a accumulations occur after marine inflow events, whereas phytoplankton does not accumulate during high water outflow, when the water residence time is short. These results suggest that changing hydrological pattern is the most important mechanism underlying phytoplankton blooms in the PLE.

A retrospective appraisal of the importance of high-resolution sampling for harmful algal blooms: Lessons from long-term phytoplankton monitoring at Sherkin Island, S.W. Ireland

The ongoing Sherkin Island Marine Station phytoplankton monitoring programme began in 1978. The present stations were established in 1980. Surveying has been carried out continuously since then, employing a more comprehensive sampling strategy than the few other published long-term records. Examples are presented here of cell concentrations for several HAB species and other bloom species from the Sherkin Island record. These show far more short-term and long-term natural variation in time and space than might be expected from the other records in the literature used to investigate HAB, suggesting that ecological complexity may be critically misrepresented in such investigations. This has serious implications for scientists evaluating reports of past HAB (what cell densities were responsible?) and predicting future HAB. Evidence presented here also suggests limitations that should be recognized in using cell concentrations from plankton in models for predictions of HAB.

Overview of Research into the Coastal Effects of the Macondo Blowout from the Coastal Waters Consortium: A GoMRI Consortium

ABSTRACT The Coastal Waters Consortium (CWC) led by Louisiana Universities Marine Consortium is one of eight Gulf of Mexico Research Initiative research consortia. The CWC focuses on: oil transport and fate, chemical evolution and biological degradation, and environmental effects.The following is an overview of a portion of the research conducted within the consortium. The consortium works in a system that was impacted by the Deepwater Horizon oil disaster and additionally impacted by freshwater diversions resulting in changes in salinity, tropical storms, and hurricanes. First, we conducted model simulations assessing oil transport into the Barataria Bay estuary, which indicate that easterly winds and feeding of the anticyclonic gyre in the Louisiana Bight pushed the oil into Barataria Bay. In subtidal sediments adjacent to oiled marshes, marsh detritus from eroding marsh edges eventually became entrained in the sediment column. Biotic impacts vary. The above-ground plant biomass appears healthy at the individual sampling sites; overall the most seaward (i.e., likely oil-impacted) areas of Terrebonne and Barataria Bay have shown, via satellite data, a distinct decline in marsh vegetation coverage since 2010. Oysters appear to be affected by predation and salinity variation. Microbial diversity from marsh-edge sediments is distinct from before and after the spill, and between unoiled and oiled marshes, with lower diversity in oiled marshes; but the greatest community composition shifts are in marshes affected by the freshwater diversions. Changes in microbial diversity in the water column at the stream-side edge of oiled marshes are extensive and are related to marsh edge erosion. In contrast, oiling of marshes had no impact on ammonia oxidizer or denitrifier abundances and on soil biogeochemical process rates 2+ years post-spill. Analysis of long-term offshore phytoplankton community and hypoxia data indicate some signal of the Macondo oil, but these components of the ecosystem remain mostly influenced by the fresh water and nutrients delivered by the Mississippi River. The consortium continues to work to tease apart oil impacts, effects of salinity, natural variation, and disturbance from tropical storms and hurricanes to determine the trajectory for health of shelf waters and Louisiana's marshes.

Revisiting the Chesapeake Bay Phytoplankton Index of Biotic Integrity

In 2006, a phytoplankton index of biotic integrity (PIBI) was published for Chesapeake Bay Lacouture et al. (Estuaries 29(4):598-616, 2006). The PIBI was developed from data collected during the first 18years (1985-2002) of the Chesapeake Bay Program long-term phytoplankton and water quality monitoring programs. Combinations of up to nine phytoplankton metrics were selected to characterize bay habitat health according to plankton community condition in spring and summer seasons across four salinity zones. The independent data available at the time for index validation was not sufficient to test the PIBI because they lacked critical index parameters (pheophytin and dissolved organic carbon) and reference samples for some seasons and salinity zones. An additional 8years of monitoring data (2003-2010) are now available to validate the original index, reassess index performance and re-examine long-term trends in PIBI conditions in the Bay. The PIBI remains sensitive to changes in nutrient and light conditions. Evaluation of the PIBI results over the entire 1985-2010 time period shows no discernible trends in the overall health of Bay habitat based on phytoplankton community conditions. This lack of overall PIBI trend appears to be a combined response to declines in water clarity and improvements in dissolved inorganic nitrogen and dissolved phosphorus conditions in the bay.

SHORT- VERSUS LONG-TERM RESPONSES TO CHANGING CO2IN A COASTAL DINOFLAGELLATE BLOOM: IMPLICATIONS FOR INTERSPECIFIC COMPETITIVE INTERACTIONS AND COMMUNITY STRUCTURE

Increasing pCO2 (partial pressure of CO2 ) in an "acidified" ocean will affect phytoplankton community structure, but manipulation experiments with assemblages briefly acclimated to simulated future conditions may not accurately predict the long-term evolutionary shifts that could affect inter-specific competitive success. We assessed community structure changes in a natural mixed dinoflagellate bloom incubated at three pCO2 levels (230, 433, and 765 ppm) in a short-term experiment (2 weeks). The four dominant species were then isolated from each treatment into clonal cultures, and maintained at all three pCO2 levels for approximately 1 year. Periodically (4, 8, and 12 months), these pCO2 -conditioned clones were recombined into artificial communities, and allowed to compete at their conditioning pCO2 level or at higher and lower levels. The dominant species in these artificial communities of CO2 -conditioned clones differed from those in the original short-term experiment, but individual species relative abundance trends across pCO2 treatments were often similar. Specific growth rates showed no strong evidence for fitness increases attributable to conditioning pCO2 level. Although pCO2 significantly structured our experimental communities, conditioning time and biotic interactions like mixotrophy also had major roles in determining competitive outcomes. New methods of carrying out extended mixed species experiments are needed to accurately predict future long-term phytoplankton community responses to changing pCO2 .

Long-term phytoplankton dynamics in a Mediterranean eutrophic lagoon (Cabras Lagoon, Italy)

The temporal variations in phytoplankton abundance and diversity were analysed from 1999 to 2009 in Cabras Lagoon, a shallow eutrophic Mediterranean lagoon (west coast of Sardinia). It is one of the aquatic ecosystems listed in the “Marine Ecosystems of Sardinia” by the Italian Network of Long-Term Ecological Research (LTER Italy). The objectives of this work were to assess the general features of phytoplankton succession and its dynamics in relation to environmental factors along the decade 1999–2009, and to identify the main forces driving phytoplankton variability in terms of the general trends in Mediterranean transitional waters. Our results highlight several important aspects: a shift in the annual peak density from the summer–autumn season, observed in 1999–2002, to the winter–spring season in subsequent years; simultaneous reductions in chlorophyll a concentrations and cell size from 1999 to 2008; and the modification of the structure of the phytoplankton assemblages, from a more differentiated class composition in 1999–2002 to the subsequent dominance of the Cyanophyceae. Moreover, analysis of similarity on both abiotic and phytoplanktonic data shows significant differences among the hydrological years considered. Pearson's correlations indicate temperature, salinity and nutrients as the environmental parameters that most influenced the phytoplankton composition and dynamics.

An analysis of long-term phytoplankton dynamics in Muskegon Lake, a Great Lakes Area of Concern

Long-term monitoring of aquatic ecosystems is essential to distinguish the effects of human-induced stressors from natural patterns of ecologic variation, especially in Great Lakes Area of Concern such as Muskegon Lake. Samples collected between 2003–2009, as part of a continuing long-term monitoring study of the lake, were analyzed to: 1) detect spatial and temporal patterns in the phytoplankton biovolume and species composition; 2) evaluate the environmental variables that regulate phytoplankton community composition changes; and 3) determine the ecological conditions under which toxin-producing Microcystis species occur. Distinct patterns in Muskegon Lake phytoplankton were not evident among sites, which can be explained, at least in part, by the well-mixed waters in this drowned river mouth system. However, surface and bottom samples within sites had significantly different biovolume and species composition, suggesting that horizontal mixing did not extend throughout the water column. Surface samples had greater phytoplankton biovolume than bottom samples, possibly because of greater irradiance. Seasonally, the least biovolume was recorded in spring samples, which were dominated by diatoms. Phytoplankton biovolume was greatest in the summer when cyanobacteria were abundant. Environmental variables that correlated highly with the ordination space defined by species composition included sulfate, specific conductance, total dissolved solids, and chloride. Results from regression tree analysis predicted increasing biovolume of Microcystis aeruginosa with increasing concentrations of the total Kjeldahl nitrogen. The seven-year period of this study did not reveal major changes in the lake's environment and phytoplankton communities, but the presence of invasive and toxin-producing species warrants their continued monitoring.

Interactions between nutrient availability and climatic fluctuations as determinants of the long-term phytoplankton community changes in Lake Garda, Northern Italy

Major focus in interpreting phytoplankton changes in specific typologies of waterbodies or in single lakes is directed towards nutrients and climatic dynamics. During the last 35 years, Lake Garda (Northern Italy; A = 368 km2, zmax = 350 m, V = 49 km3) underwent a significant increase of phosphorus in the water column, from ca. 10 μg P l−1 to 18–22 μg P l−1. At the multi-decadal scale, the increase of the trophic status had a positive impact on the growth of Cyanobacteria (mainly Oscillatoriales) and, partly, diatoms, as demonstrated by the long-term ecological research carried out since the beginning of the 1990s in the deepest zone of the lake. Conversely, the increase of Peridiniales (mostly Ceratium hirundinella) in the recent years appeared also associated with the interannual variations of lake temperature. At the seasonal and annual scale, the development of the large diatoms and Oscillatoriales during the periods of their maximum growth (early spring, and summer and autumn, respectively) was strongly controlled by the extent of spring vertical water mixing and nutrient fertilization of surface waters, which, in turn, were negatively dependent on the air and water temperatures in winter and early spring. Therefore, contrary to the positive impact of milder winters on phytoplankton growth in many lakes of high latitudes, warmer winter temperatures in deep oligomictic lakes of lower latitudes can determine periodic shifts towards more oligotrophic conditions and a minor development of diatoms and specific harmful cyanobacterial groups (Oscillatoriales). The complex relationships between the explanatory and response variables were tested by applying Path Analysis (Structural Equation Modeling). This multiequational technique has great potential for studying causal relationships in temporally ordered variables. The results highlight the necessity to study the consequences of climatic fluctuations on the phytoplankton communities at different temporal scales and complexity, also including the indirect effects of climatic dynamics mediated by the morphometric, morphological and hydrological characteristics of lakes, and the possible synergic or opposite effects with other forcing variables, including nutrients.

Long‐term phytoplankton community changes in a deep subalpine lake: responses to nutrient availability and climatic fluctuations

Summary1. In natural lakes, modifications in the species composition and abundance of phytoplankton communities may ultimately be responses to changes in nutrient availability and climatic fluctuations. Phytoplankton and associated environmental factors were collected at monthly intervals from the beginning of the 1990s to 2007 in the large subalpine Lake Garda (zmax = 350 m, V = 49 × 109 m3). In this study period, the lake showed a slight and continuous increase of total phosphorus (TP) in the water column, up to concentrations of 18–20 μg P L−1. This increase represented the last stage of a long‐term process of enrichment documented since the 1970s, when concentrations of TP were below or around 10 μg P L−1.2. At the community level, annual phytoplankton cycles underwent a unidirectional and slow shift mainly due to changes in the species more affected by the nutrient enrichment of the lake. After a first and long period of dominance by conjugatophytes (Mougeotia) and diatoms (Fragilaria), phytoplankton biomass in recent years was sustained by cyanobacteria (Planktothrix). Other important modifications in the development of phytoplankton were superimposed on this pattern due to the effects of annual climate fluctuations principally mediated by the deep mixing events at spring overturn and, secondarily, by temperature and thermal stability of the water column during the growing season.3. Interannual variations in the stability and temperature of the water column appeared to influence the development of a few subdominant flagellates (dinophytes and cryptophytes). Nevertheless, the major impact of climate on phytoplankton was indirect, and mediated through the effects of winter climatic conditions on deep mixing dynamics. Winter climatic fluctuations proved to be a key element in a linked chain of causal factors including cooling of hypolimnetic waters, deep vertical mixing and epilimnetic nutrient replenishment. The process of fertilisation was measurable both for TP and dissolved inorganic nitrogen, although only the first had a large effect, reinforcing the seasonal growth of a few dominant groups. The degree of nutrient replenishment further increased the spring development of large diatoms and the increase of Planktothrix in summer and autumn.4. Currently, changes in nutrient concentrations have the greatest effect on the phytoplankton community, while direct effects due to the interannual variations in the thermal regime are of secondary importance compared with the indirect effects mediated through deep water mixing and spring fertilisation. Overall, the results demonstrate that the consequences of climatic fluctuations and climate warming on phytoplankton communities need to be studied at different levels of complexity and integration, from the direct effects of temperature and thermal regime, to the indirect effects mediated by the physiographic characteristics of water bodies.