Did the evolution of the phytoplankton fuel the diversification of the marine biosphere?

We discuss the possible links between the fossil record of marine biodiversity, nutrient availability and primary productivity. The parallelism of the fossil records of marine phytoplankton and faunal biodiversity implicates the quantity (primary productivity) and quality (stoichiometry) of phytoplankton as being critical to the diversification of the marine biosphere through the Phanerozoic. The relatively subdued marine biodiversity of the Palaeozoic corresponds to a time of relatively low macronutrient availability and poor food quality of the phytoplankton as opposed to the diversification of the Modern Fauna through the Mesozoic–Cenozoic. Increasing nutrient runoff to the oceans through the Phanerozoic resulted from orogeny, the emplacement of Large Igneous Provinces (LIPs), the evolution of deep-rooting forests and the appearance of more easily decomposable terrestrial organic matter that enhanced weathering. Positive feedback by bioturbation of an expanding benthos played a critical role in evolving biogeochemical cycles by linking the oxidation of dead organic matter and the recycling of nutrients back to the water column where they could be re-utilized. We assess our conclusions against a recently published biogeochemical model for geological time-scales. Major peaks of marine diversity often occur near rising or peak fluxes of silica, phosphorus and dissolved reactive oceanic phosphorus; either major or minor ⁸⁷Sr/⁸⁶Sr peaks; and frequently in the vicinity of major (Circum-Atlantic Magmatic Province) and minor volcanic events, some of which are associated with Oceanic Anoxic Events. These processes appear to be scale-dependent in that they lie on a continuum between biodiversification on macroevolutionary scales of geological time and mass extinction.     

Rarity,ecological memory,rate of floral change in phytoplankton—and the mystery of the Red Cock

In this article, we attempt to estimate the contemporary phytoplankton species pool of a particular lake, by assessing the rate of floral change over a period of 15 years. Phytoplankton time series data from Lake Stechlin, an oligo-mesotrophic lake in the Baltic Lake District (Germany) were used. Of the 254 algal species recorded during the 15-year of studies with roughly biweekly sampling, 212 species were planktonic. In the individual plankton years, the recorded total number of species changed between 97 and 122, of which the number of dominants (>1% contribution to the annual average of total biomass) was only 10–19. The 15-year cumulative number of species exhibited an almost linear increase after an initial saturation phase. This increase was attributed to two reasons: increase of sample size and immigration of species new to the flora. Based on a probabilistic model developed in this study, we estimated the number of co-existing planktonic species of the lake as some 180, and the rate of floral change as 1–2 species per year. Of these co-existing species, only few maintain the matter–energy processing ecosystem functions in any particular plankton year. Selection of these dominants is probably driven by mesoclimatic cycles, coupled with human-induced forcing, like eutrophication. All others are hiding as an ecological memory, in the sense of the capacity or experiences of past states to influence present or future responses of the community. Data analyses suggest that selection of the ‘memory species’ that show temporary abundance increases over shorter (several years) periods are largely dependent upon the dominants. These results show that interspecific interactions and the particular autecological features of the dominants, together with their effects on the whole ecosystem, act as a major organizing force. Some phytoplankton species, like Planktothrix rubescens, are efficient ecosystem engineers with cascading effects of both a top-down and bottom-up nature. Historical scientific data on Planktothrix blooms in Lake Stechlin suggest cyclic patterns in long-term development of phytoplankton which, as the legend of the Red Cock suggests, dates back much further than scientific archives.     

Seasonal succession of phytoplankton in a lake of the Paraná river floodplain,Argentina

Phytoplankton biomass, morphological and taxonomic composition, species diversity and productivity were analyzed in a shallow lake of the Middle Paraná River floodplain (El Tigre, 31 ° 41 S and 60° 42 W), between November 1986 and July 1988. Lake inundation (filling and through-flow phases) constituted an intense long-term perturbation in the physical and chemical environment. As the lake filled with river water, K-selected species (netplanktonic filamentous bluegreens, > 37 µm, with low surface area/volume (SA/V) ratios) that had existed prior to filling (late spring 1986) were replaced in summer-fall by r-selected species (nannoplanktonic chlorophytes and cryptophytes, < 37 µm, mainly stout forms with high SA/V ratios). During the through-flow phase, lentic phytoplankton was replaced by lotic flagellate populations due to the direct flushing by river water. During the period of falling water (drainage and isolation phases), nanoplanktonic algae with similar characteristics to those of the filling phase dominated in late winter-spring. Later in the isolation phase, these were succeeded by K-selected species (netplanktonic algae, mainly motile spherical dinoflagellates and filamentous bluegreens with low SA/V ratios). Simultaneously, primary production per unit biomass decreased and total biomass and specific diversity increased. Seasonal changes of phytoplankton in floodplain lakes can be interpreted as the interaction between true successional development (as observed in the drainage and isolation phases) and intermediate disturbance. Using Reynolds' terminology, short-term disturbance (slight inflow of nutrient-rich river water) caused reversion to an earlier stage in the former succession, and long-term disturbance (lake inundation) truncated the successional progression and a new (or shifted) succession was initiated.     

Temporal fluctuations of the phytoplankton community of the Baía River, in the upper Paraná River floodplain, Mato Grosso do Sul, Brazil

Subsurface samples were taken monthly (March/1993 through February/1994) in the Baía River, a tributary of the right bank of the Paraná River (22° 43 S and 53° 17 W). We analyzed temporal changes in the phytoplankton community in relation to density, biomass, richness and species diversity, equitability, size structure, and dominance. We related these to regional climate and hydrology, and to the physical and chemical variables of the water column. We determined 119 taxa, wich were numerically dominated by the class Chlorophyceae with 37 taxa. The classes Cyanophyceae and Bacillariophyceae contributed the most abundant biomass and were responsible for the two peaks that were observed. The high water period was generally characterized by lower phytoplankton biomass, higher richness and species diversity, and higher density of nannoplanktonic species such as Monoraphidium tortile in March, and Cryptomonas brasiliensis in January. During low water, however, the highest values of phytoplankton biomass were recorded, with heterocytous cyanophytes dominating during the phase of greater stability of the water column, and filamentous diatoms during periods of mixing turbulence. The fluctuations observed indicate that the hydrosedimentological regime of the Paraná River, together with the climatological factors, constitute the main forcing functions acting on the Baía River phytoplankton.     

Horizontal mesoscale distribution of phytoplankton in the Tvärminne sea area,southern Finland

Horizontal distribution of phytoplankton was studied by sampling two grids, both having 16 sampling points. The distance between sampling points was 10 m. Phytoplankton species were enumerated by inverted microscopy and the observed distribution patterns were compared with a random distribution. Contagious distribution was common among the species studied. Both actively moving and passively drifting species showed significant contagiousness in their distribution. Actively moving and passively drifting species showed different distribution patterns. The effects of horizontal heterogeneity on phytoplankton sampling are discussed.     

Studies on the phytoplankton of the River Ganges,Varanasi, India,Part II “The seasonal growth and succession of the plankton algae in the River Ganges”

Summary An attempt has been made to find out the productivity and periodicity of the phytoplankton in relation to the physical and chemical characteristics of the river Ganges, between University Ghat and Raj Ghat at Varanasi. The observations were taken during a single year (i.e., March 1957 to March 1958). The chemical analyses of the water and estimates of the number of phytoplankton were made on samples collected fortnightly.The phytoplankton has proved to be eutrophic and polymixic in nature. Diatoms formed the largest bulk, blue-greens being the second major constituent while the green algae formed the third in order of abundance with, however, large number of species. A definite phytoplankton periodicity has been noticed and found to be seasonal in character.Correlations between the phytoplankton periodicity and population maxima, with the habitat factors revealed depletion of nutrients like, nitrogen, phosphate, silica, carbon, magnesium and sulphates.Changes in the phytoplankton populations were clearly evident more in relation to physical than to chemical conditions of the water. Changes in water-level, transparency and temperature affected the growth of the phytoplankton.High alkalinity and buffering capacity resulted in preferential growth of diatoms and blue-green algae. Rich phosphates and silicates coupled with moderate nitrogen contents were responsible for high phytoplankton yields in summer and winter seasons.The abundance of blue-greens may be due to the higher values of pH, temperature, dissolved organic matter, phosphate, nitrogen, and relatively high values of dissolved oxygen. Green algae showed a wide range of adaptability but could not develop in any abundance.     

Lack of pattern among phytoplankton assemblages. Or,what does the exception to the rule mean?

In Science we cannot say that `the exception proves the rule'. We have been looking to define patterns in phytoplankton occurrence across trophic spectra where conspicuous covariations between algae and trophic states have been reported. We consider quite different phytoplankton communities observed under similar trophic conditions: we illustrate this point by considering five different phytoplankton communities living in five water bodies in the same wetland, along a TP gradient and over a period of 2 years. This system showed a remarkable dissimilarity of species representation, implying communities of uncorrelated species vary considerably over time. Despite the presence of some characteristic species, communities were not related to a given trophic state. However, coarser community attributes, such as clusters of taxonomic classes, appeared to be more useful in identifying patterns and assembly rules related to trophic spectra. Some ecological concepts can be related to this lack of pattern, e.g., nonconvergence, trajectories far from equilibrium and assembly rules of communities.     

A principal components analysis of the phytoplankton from a pond in the Paraná River Valley,Argentina

A principal-component scheme was applied to a set of 10 physical, chemical, and biological attributes, measured in 42 samples collected from Los Matadores pond (May 1974–July 1975). The first three components account for 82.1% of the total variance in the data and represent the lotic influence (58.3% of the variance), the seasonal cycle (16.4%), and the degree of maturity (7.4%).The authors are members of the Carrera del Investigador Cientifico y Tecnologico, Consejo Nacional de Investigaciones Cientificas y Tecnicas (CONICET), Argentina.     

Survival strategies of some species of the phytoplankton community in the Barra Bonita Reservoir (São Paulo,Brazil)

The dynamics of the phytoplankton community in the Barra Bonita Reservoir (S?o Paulo, Brazil) were studied through daily sampling in the field (integrated samples from the euphotic zone) and microcosm experiments, for two short periods: the winter of 1993 (June 30 to July 10) and the summer of 1994 (January 24 to February 2). The goal of the study was to evaluate and compare the variations in the composition of isolated phytoplankton community which occur over short periods of time. Three series were separated into Erlenmeyer flasks for each study period, with samples from the euphotic zone divided into three portions: total, smaller than 64 μm, and smaller than 20 μm. All of the Erlenmeyer flasks were inclubated in situ at the sampling station. The maximum period of incubation was 10 d. Variations of the community in the euphotic zone were characterised by high diversity and a community in a state of non-equilibrium in winter, without the predominance of any species. In the summer, the community presented a low diversity and a state of equilibrium, with the predominance of Microcystis aeruginosa. The microcosm experiments showed that the confinement of the community in the Erlenmeyer flasks, and therefore in isolation from the physical variability of the ecosystem, especially in relation to the mixing patterns, stimulated the return of the community to the initial phases of succession with the predominance of small species and those which grow rapidly (r-selective or C-strategist). This revised version was published online in July 2006 with corrections to the Cover Date.     

Peculiarities of the state of phytoplankton in the Baltic Sea’s Vistula Lagoon at the turn of 21st century

In 1989–2005 the qualitative composition and seasonal and inter-annual dynamics of the total biomass of phytoplankton, chlorophyll a content, hydrological, and hydrochemical parameters were studied in the Russian zone of the Vistula Lagoon in the Baltic Sea. The results of these observations were compared to the data obtained in the 1950s and 1970s. The structural reorganization of phytoplankton was revealed, testifying to the negative changes in the ecosystem under conditions of anthropogenic pollution and eutrophication, climate warming, and increasing salinity.     

Why are there about 5000 species of phytoplankton in the sea?

The relative abundances of phytoplankton taxa conform approximatelyto a finite geometric series in which there are 20–25species per decade of ranked abundance. Such series can contain160–400 species between the commonest (10²²–10²⁶cells) and the rarest (10¹⁰–10¹⁴ cells). Thus, between12 and 31 such series are needed to explain the observed diversity,5x10³ species, of marine phytoplankton. The number of seriesis similar to the number (20–25) of upper-ocean watermasses defined by dilution time scale of order 10¹–10²years. Interpretations of this coincidence are discussed.     

Downstream variations of phytoplankton in the St.Lawrence River (Québec, Canada)

Longitudinal variations of phytoplankton biomass and composition were assessed in a 250 km-long section of the St.Lawrence River, which alternately runs through narrow (< 2 km) river cross sections and wide (up to 10 km) fluvial lakes. In the main river stem, concentrations of suspended matter and total phosphorus increased with distance downstream, whereas light penetration decreased. Seasonal changes in plankton composition and biomass were more important than those resulting from differences in water mass (tributary) of origin. Sampling at three cross river sections and in two fluvial lakes showed a progressive downstream decrease in phytoplankton biomass and changes in size structure and taxonomic composition. River plankton was primarily composed of small (< 10 µm equivalent spherical diameter), truly planktonic cells belonging to Cryptophyceae and diatoms, with Chlorophyceae in summer. Plankton sampled in summer among rooted macrophytes in fluvial lakes exhibited a higher biomass of resuspended periphytic algae than in the main river stem, which contributed slightly to downstream phytoplankton biomass.Successive river cross sections always shared about 50% of their taxa, indicating a rapid downstream transport of algae within the main water mass. However, the proportion of species common to all cross sections was highest during the spring freshet, and lowest during summer low discharge, likely resulting from the development of a distinct flora in fluvial lakes during summer. Conversely, about 30% of the identified taxa were exclusive to a cross section and were replaced by others occurring downstream. Overall, phytoplankton composition along the St.Lawrence River is primarily controlled by advective forces, which result in a homogeneous flora in the main river stem, with a local contribution of resuspended periphyton from fluvial lakes.     

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.     

Are copepods important grazers of the spring phytoplankton bloom in the western Irish Sea?

The spring phytoplankton bloom and copepod grazing were studied at a coastal and offshore station in the western Irish Sea during 1997. Maximum chlorophyll standing stocks of 132.8 mg m^-2 inshore and 199.4 mg m^-2 offshore were measured in late April. At that time, mean water column temperatures were 10 and 8C at the coastal and offshore station, respectively. Spring bloom production at the coastal station was estimated as 31.2 g C m^-2 and was dominated by the diatom Guinardia delicatula. Offshore, production was 28.2 g C m^-2 and the bloom was composed of small (10 m) phytoflagellates and the silicoflagellate Dictyocha speculum. Maximum copepod abundance (189 and 544 x 10³ individuals m^-2, inshore and offshore, respectively) coincided with the spring bloom. Pseudocalanus and Temora ingestion rates were derived from measurements of gut pigment fluorescence, and were found to vary during the course of the spring bloom as a result of changes in gut content. Grazing by late copepodite and adult Pseudocalanus and Temora was variable inshore, but overall accounted for 17% of bloom production. Offshore, 22% of bloom production was grazed with maximum grazing (76% of daily production) occurring at the end of the bloom. Large copepod species were not major grazers of the spring bloom. Greater utilization of spring bloom production by copepods in the western Irish Sea compared to regions of the North Sea is attributed to differences in population size at the time of the bloom.      

Erratum to: How do UV radiation, temperature, and zooplankton influence the dynamics of alpine phytoplankton communities?

Plankton in mountain lakes are confronted with generally higher levels of incident ultraviolet radiation (UVR), lower temperatures, and shorter growing seasons than their lower elevation counterparts. The direct inhibitory effects of high UVR and low temperatures on montane phytoplankton are widely recognized. Yet little is known about the indirect effects of these two abiotic factors on phytoplankton, and more specifically whether they alter zooplankton grazing rates which may in turn influence phytoplankton. Here, we report the results of field microcosm experiments that examine the impact of temperature and UVR on phytoplankton growth rates and zooplankton grazing rates (by adult female calanoid copepods). We also examine consequent changes in the absolute and relative abundance of the four dominant phytoplankton species present in the source lake (Asterionella formosa, Dinobryon sp., Discostella stelligera, and Fragilaria crotonensis). All four species exhibited higher growth rates at higher temperatures and three of the four species (all except Dinobryon) exhibited lower growth rates in the presence of UVR versus when shielded from UVR. The in situ grazing rates of zooplankton had significant effects on all species except Asterionella. Lower temperatures significantly reduced grazing rates on Fragilaria and Discostella, but not Dinobryon. While UVR had no effect on zooplankton grazing on any of the four species, there was a significant interaction effect of temperature and UVR on zooplankton grazing on Dinobryon. Discostella and Dinobryon increased in abundance relative to the other species in the presence of UVR. Colder temperatures, the presence of zooplankton, and UVR all had consistently negative effects on rates of increase in overall phytoplankton biomass. These results demonstrate the importance of indirect as well as direct effects of climate forcing by UVR and temperature on phytoplankton community composition in mountain lakes, and suggest that warmer climates and higher UVR levels may favor certain species over others.     

Wetland macrophyte diversity, trophic status and phytoplankton： a case study of the cascading effects of an invasive herbivore

Shallow water bodies can exist in alternative stable states, a clear water state with high coverage of macrophytes or a turbid state with high phytoplankton biomass. The alternative equilibria hypothesis has been proposed to explain the occurrence of the alternative stable states (Scheffer et al., 1993)[1], which assumes that: 1),     

Hydrologic and climatic regimes limit phytoplankton biomass in reservoirs of the Upper Paraná River Basin,Brazil

In reservoirs of the Upper Paraná River Basin, Brazil Paraguay, phytoplankton biomass is generally low relative to reservoirs in other parts of the world. To investigate what might be limiting phytoplankton biomass in the Upper Paraná reservoirs, we used an empirical approach wherein climatic conditions in the basin as well as chemical, hydrological and morphometrical characteristics of 13 reservoirs, were compared against those prevailing in 58 reservoirs of the south-central United States. In both regions, phytoplankton biomass was correlated with chemical, hydrological and morphometrical characteristics of the reservoirs. There were no differences in phosphorus concentrations between the two regions, nitrogen levels were higher in the Upper Paraná, ionic concentrations were higher in the south-central United States and there were major differences in all physical characteristics considered. In reservoirs of the south-central United States, hydraulic retention time increased and discharges decreased during the post-spawning period allowing increases in phytoplankton biomass; in the Upper Paraná River Basin, discharges peaked during the post-spawning period, delaying the peak of phytoplankton blooms until after completion of the post-spawning period. Our comparison between the two sets of reservoirs suggests that hydrologic regimes dictate differences in phytoplankton biomass and that hydrology of the Upper Paraná River Basin, exacerbated by climatic patterns, deter phytoplankton production despite suitable levels of essential nutrients.     

Eutrophication of the salt valley reservoirs, 1968–73 I. The effects of eutrophication on standing crop and composition of phytoplankton

A study was carried out over a period of six years to determine the effects of eutrophication upon standing crop and composition of the phytoplankton in four recently constructed flood control reservoirs in Nebraska. Water samples collected weekly during June, July, and August from 1968–73 were analyzed for chlorophyll a, phytoplankton composition, and phytoplankton abundance. Total volume of phytoplankton was calculated from appropriate dimensions and formulae. Inorganic turbidity in one reservoir was an important factor regulating the size and composition of the phytoplankton standing crop. In that reservoir diatoms were the most important component of the phytoplankton community during those years in which inorganic turbidity was greatest. When inorganic turbidity declined, blue-green algae became dominant.In the clear-water reservoirs chlorophyll a, phytoplankton number, and phytoplankton volume were significantly correlated with reservoir age, with the oldest containing chlorophyll a concentrations up to 247 mg/m³ and mean phytoplankton volumes up to 329 mm³/l, values sufficient to place it in the hypereutrophic category. The reservoirs had 2–6 times more chlorophyll a present at the end of the study than at the beginning, with the increase being greatest in the newest reservoir. Phytoplankton volume was significantly correlated with chlorophyll a in all the reservoirs.Blue-green algae quickly became established as community dominants in the reservoirs, making up over 80 percent of the phytoplankton volume in the newest reservoir by the second year of its existence. In the other clear-water reservoirs, blue-greens usually constituted over 95 percent of the total phytoplankton volume in summer. three genera, Microcystis, Aphanizomenon, and Anabaena, were responsible for virtually the entire standing crop of blue-greens in all the reservoirs.     

Did sulfate availability facilitate the evolutionary expansion of chlorophyll a+c phytoplankton in the oceans?

During the Mesozoic Era, dinoflagellates, coccolithophorids and diatoms became prominent primary producers in the oceans, succeeding an earlier biota in which green algae and cyanobacteria had been proportionally more abundant. This transition occurred during an interval marked by increased sulfate concentration in seawater. To test whether increasing sulfate availability facilitated the evolutionary transition in marine phytoplankton, the cyanobacterium Synechococcus sp., the green alga Tetraselmis suecica and three algae containing chlorophyll a+c (the diatom Thalassiosira weissflogii, the dinoflagellate Protoceratium reticulatum and the coccolithophorid Emiliania huxleyi) were grown in media containing 1, 5, 10, 20, or 30 mm SO(4) (2-) . The cyanobacterium and the green alga showed no growth response to varying [SO(4) (2-) ]. By contrast, the three chlorophyll a+c algae showed improved growth with higher [SO(4) (2-) ], but only up to 10 mm. The chlorophyll a+c algae, but not the green alga or cyanobacterium, also showed lower C:S with higher [SO(4) (2-) ]. When the same experiment was repeated in the presence of a ciliate predator (Euplotes sp.), T. suecica and T. weissflogii increased their specific growth rate in most treatments, whereas the growth rate of Synechococcus sp. was not affected or decreased in the presence of grazers. In a third experiment, T. suecica, T. weissflogii, P. reticulatum and Synechococcus sp. were grown in conditions approximating modern, earlier Paleozoic and Proterozoic seawater. In these treatments, sulfate availability, nitrogen source, metal availability and Pco(2) varied. Monospecific cultures exhibited their highest growth rates in the Proterozoic treatment. In mixed culture, T. weissflogii outgrew other species in modern seawater and T.suecica outgrew the others in Paleozoic water. Synechococcus sp. grew best in Proterozoic seawater, but did not outgrow eukaryotic species in any treatment. Collectively, our results suggest that secular increase in seawater [SO(4) (2-) ] may have facilitated the evolutionary expansion of chlorophyll a+c phytoplankton, but probably not to the exclusion of other biological and environmental factors.