Picophytoplankton, nanophytoplankton, heterotrohpic bacteria and viruses in the Changjiang Estuary and adjacent coastal waters

Flow cytometry (FCM) was used to examine the abundances anddistributions of different picophytoplankton groups (i.e. Synechococcus,Prochlorococcus and picoeukaryotes), nanophytoplankton, heterotrophicbacteria and viruses were examined in the Changjiang Estuary,China and adjacent coastal waters during autumn 2004. Watertemperature and light availability were found to be criticalfactors for picophytoplankton growth. Positive correlationswere found between picophytoplankton, heterotrophic bacteriaand viruses, and a seaward-increasing trend in the V-I (thegroup yielding high green fluorescence according to FCM) populationwithin viruses was detected. The importance of nanophytoplanktonis progressively usurped by picophytoplankton with increasingdistance offshore. Picoeukaryotes are the most successful groupamong picophytoplankton in near-shore eutrophic waters, whereasProchlorococcus surpasses other groups within the pico- andnanophytoplankton community in offshore oligotrophic regionsof the East China Sea Shelf.     

Dynamics of prokaryotic picoplankton community in the central and southern Adriatic Sea (Croatia)

This paper addresses the dynamics of the prokaryotic picoplankton community in the coastal and open sea areas of the central Adriatic and in the coastal area of the southern Adriatic. This involved the study, from January to December 2005, of bacteria (total number of non-pigmented bacteria; high nucleic acid content (HNA) bacteria; low nucleic acid content (LNA) bacteria), cyanobacteria (Synechococcus and Prochlorococcus) and heterotrophic nanoflagellates. During the warmer seasons, in the mainly oligotrophic area under investigation into the Adriatic Sea, bacterial densities and bacterial production have shown an increase in values and domination of the LNA group of the bacterial population. In contrast, in those areas influenced by karstic rivers, the domination of HNA bacteria in total abundance of non-pigmented bacteria and high values of bacterial production was estimated throughout the investigated period. Our results show the importance of both HNA and LNA bacterial groups in the total bacterial activity throughout the investigated area. The biomass of bacteria was mostly predominant in the prokaryotic community, while within the autotrophic community Synechococcus biomass mostly predominated. During the warmer seasons, an increase in autotrophic biomass was observed in relation to non-pigmented biomass. The importance of predation in controlling bacteria by heterotrophic nanoflagellates was pronounced during the warmer period and in the coastal areas.     

Picoplankton community structure along the northern Iberian continental margin in late winter-early spring

The surface distribution of autotrophic and heterotrophic picoplanktonwas assessed in 24 transects perpendicular to the coast alongthe N and NW Iberian peninsula shelf in late winter and earlyspring 2002. Community structure was analyzed by flow cytometry(FC) and found to be strongly influenced by hydrography. Typicallate winter conditions were found during the survey, characterizedby the presence of the poleward Portugal coastal counter current(PCCC) in the west and an increasing stratification eastwards.Cyanobacteria (mostly Synechococcus) dominated at low chlorophylla (Chl a) concentration whereas both the total and relativeabundance of picoeukaryotes generally increased with total phytoplanktonbiomass. Differences in the cell size of most FC-defined picoplanktonicgroups were also observed along the longitudinal and coastal–offshoregradients. The presence of Prochlorococcus (<10³ cells mL^–1)coincided with the core of the PCCC and its significant correlationwith salinity suggests its possible use as a tracer of thiscurrent. Two groups of heterotrophic bacteria were distinguishedaccording to their relative DNA content. High DNA bacteria dominatedthe community (60 ± 1% SE of total numbers), reachingmaximum values in areas under riverine influence with presumedhigher inputs of organic matter. Picoplankton biomass was dominatedby heterotrophic bacteria in the western region (58 ±3%) while autotrophic groups contributed on average 66 ±2% in the southern Bay of Biscay. The heterotrophic bacteriato phytoplankton biomass ratio decreased significantly alongthe measured range. Yet showing regional differences, the estimatedcontribution of picophytoplankton to total algal biomass washigh (mean 59 ± 4%), indicating the important role ofsmall cells at the onset of the spring bloom in these temperateshelf waters.     

Distribution of microbial populations and their relationship with environmental parameters in the coastal waters of Qingdao,China

In order to understand the large‐scale distribution of microbial populations simultaneously and their relationship with environmental parameters, flow cytometry was used to analyse samples collected from 46 stations in the coastal waters of Qingdao in spring, 2007. The distribution of virus was significantly and positively correlated with heterotrophic bacteria. Two groups of picophytoplankton (Synechococcus and picoeukaryotes) were detected; however, Prochlorococcus was not found. Picoeukaryotes and nanophytoplankton were abundant in the near‐shore waters, whereas Synechococcus was abundant in the off‐shore areas. No variation was found in vertical distribution of virus, heterotrophic bacteria, Synechococcus and nanophytoplankton abundances, except picoeukaryotes abundance in the bottom layer was dramatically lower than that in the upper layers. Correlation analyses indicated that the relationship between abiotic variables and heterotrophic bacteria, pico‐ and nanophytoplankton was closer than that between abiotic variables and virioplankton. Temperature and nutrients were the synchronous factors controlling the growth of heterotrophic bacteria, pico‐ and nanophytoplankton in the coastal waters of Qingdao in spring. The results suggested that synergistic and antagonistic effects existed among microbial groups.     

Effects of UV radiation on grazing by two marine heterotrophic nanoflagellates on autotrophic picoplankton

Phytoplankton <4-3 µm in diameter, or autotrophic picoplankton,can constitute the majority of the biomass and productivityof photosynthetic organisms in marine and freshwater systems.Indirect evidence has indicated that mortality of autotrophicpicoplankton occurs principally at night in the open ocean,but continuously in coastal water. The predominant view of thefate of autotrophic picoplankton production in the ocean isthat they are consumed by heterotrophic nanoflagellates. A possiblemechanism to explain these observations is that grazing of heterotrophicnanoflagellates on autotrophic picoplankton is inhibited byultraviolet radiation (W), at least in clear open-ocean environments.A series of laboratory experiments was conducted to examinethe effects of UV radiation on the grazing impact of two heterotrophicnanoflagellates on Synechococcus spp., a commonly occurringgenus of autotrophic picoplankton. The two nanoflagellates usedwere Paraphysomonas bandaieensis and Paraphysomonas imperforata.For both nanoflagellates, there was an inverse relationshipbetween the grazing mortality of Synechococcus and UV irradiance.The grazing mortality of Synechococcus was reduced less withP.imperforata than with P.bandaiensis. In some experiments,the effect of UV on the grazing impact of the nanoflagellatepopulations was caused in part by UV-related reductions in nanoflagellatesurvival. However, UV reduced the grazing impact of nanoflagellatesprimarily by reducing the rates of consumption of Synechococcusby individual nanoflagellates, to a degree directly relatedto UV irradiance. The results suggat that UV radiation may bean important selection factor in clear open-ocean water, andthat in order to predict the effect of increasing UV radiationon marine microbial plankton communities, we must consider interactionsbetween trophic levels as well as effects on single trophiclevels.     

Productivity, photosynthetic pigments and hydrology in the coastal front of the Eastern English Channel

In the French area of the Eastern English Channel, the coastalwaters 3 or 4 miles wide are separated from offshore watersby a coastal front. The phytoplanktonic biomass (B) and production(P) are higher in coastal waters, and an increase of biomasshas been observed at the interface. This frontal biomass, lessproductive, resulted from hydrodynamic accumulation, not dueto local growth. The productivity (P/B) deficit (50%) spreadsover the external area of the front, probably because the biomassis partly constituted of exported inactive coastal phytoplankton.The productivity is higher again in offshore waters. The variationof the pigment pool indicates a stress on phytoplankton in thefront and this ergocline was not productive at the time of thetwo cruises.     

Diel abundance and productivity patterns of autotrophic picoplankton in the lower Chesapeake Bay

Diet studies associating phototrophic picoplankton abundanceand productivity to tidal flow and pycnocline formation wereconducted during summer and winter. Higher picoplankton concentrationsand productivity were associated with waters above the pycnoclinein summer, with picoplankton abundance highest during ebb tidewhen phycocyanin enriched Synechococcus sp. were dominant. Inwinter, greater concentrations and productivity occurred inbottom waters and during flood tide, when phycoerythrin enrichedSynechococcus sp. were dominant.     

Contrasting effects of substrate and grazer manipulations on picoplankton in oceanic and coastal waters off Brazil

In two contrasting regions off the coast of Brazil, picoplankton(<1 µm) responses to removal of larger grazers andto the additions of glucose and amino acids were determined.Effects of glucose and amino acid additions (1 µM) onparticulate nitrogen and chlorophyll a concentrations, and onrates of NH₄⁺ uptake and regeneration, were observed after 5h pre-incubation. In the oceanic waters, removal of the >1µm fraction had no significant effect on the chlorophylla of the picoplankton after 5 h. However, the addition of glucosestimulated both uptake and regeneration by a mean of 27%, andthe addition of arginine led to significant decreases in therates of NH₄⁺ uptake and regeneration. In contrast, in the coastalwaters, significant increases in chlorophyll and particulatenitrogen concentrations were found after 5 h incubation in boththe amended samples and in the controls, and mean rates of NH₄⁺uptake and regeneration were affected to a lesser degree bythe additions of either glucose or amino acids than in the oceanicwaters. The oceanic responses were suggestive of carbon limitationof heterotrophic bacteria. In the coastal region, on the otherhand, the supply of organic carbon and nitrogen was likely tohave been sufficient to meet the nutritional requirements ofthe heterotrophic bacteria and cyanobacteria. Grazing by largerorganisms on the picoplankton appeared to play a more significantrole in the nitrogen cycle in the coastal waters than in theoceanic waters.     

Decrease in the autotrophic-to-heterotrophic biomass ratio of picoplankton in oligotrophic marine waters due to bottle enclosure

We investigated the effects of bottle enclosure on autotrophic and heterotrophic picoplankton in North and South subtropical Atlantic oligotrophic waters, where the biomass and metabolism of the microbial community are dominated by the picoplankton size class. We measured changes in both autotrophic (Prochlorococcus, Synechococcus, and picoeukaryotes) and heterotrophic picoplankton biomass during three time series experiments and in 16 endpoint experiments over 24 h in light and dark treatments. Our results showed a divergent effect of bottle incubation on the autotrophic and heterotrophic components of the picoplankton community. The biomass of picophytoplankton showed, on average, a >50% decrease, mostly affecting the picoeukaryotes and, to a lesser extent, Prochlorococcus. In contrast, the biomass of heterotrophic bacteria remained constant or increased during the incubations. We also sampled 10 stations during a Lagrangian study in the North Atlantic subtropical gyre, which enabled us to compare the observed changes in the auto- to heterotrophic picoplankton biomass ratio (AB:HB ratio) inside the incubation bottles with those taking place in situ. While the AB:HB ratio in situ remained fairly constant during the Lagrangian study, it decreased significantly during the 24 h of incubation experiments. Thus, the rapid biomass changes observed in the incubations are artifacts resulting from bottle confinement and do not take place in natural conditions. Our results suggest that short (<1 day) bottle incubations in oligotrophic waters may lead to biased estimates of the microbial metabolic balance by underestimating primary production and/or overestimating bacterial respiration.     

The distributions of autumn picoplankton in relation to environmental factors in the reservoirs along the Wujiang River in Guizhou Province,SW China

Autumn picoplankton (Synechococcus, picoeukaryotes and heterotrophic bacteria) and environmental factors have been investigated in a series of reservoirs along the Wujiang River in Guizhou Province, SW China. The average abundances of Synechococcus, picoeukaryotes and heterotrophic bacteria was 10⁴, 10² and 10⁶ cells ml⁻¹, respectively. In autumn meso-eutrophic reservoirs, thermal stratification was clear and abundances of different picoplankton groups in release water was low; whereas these phenomena were not obvious in autumn hypereutrophic reservoir. Picoplankton numbers decreased with increasing water depth and showed a positive correlation with water temperature, which reflected the importance of light and temperature on the picoplankton growth. Contribution of Synechococcus to total phytoplankton production and contribution of picoeukaryotes to total phytoplankton production asynchronous changed with varying trophic states. Synechococcus preferred meso-eutrophic reservoirs over hypereutrophic reservoir and picoeukaryotes showed no preference for the investigated reservoirs in autumn. Handling editor: L. Naselli-Flores     

On-board flow cytometric observation of picoplankton community structure in the East China Sea during the fall of different years

On-board flow cytometric determinations of picoplankton abundance (i.e. Synechococcus spp., Prochlorococcus spp., picoeukaryotes and also heterotrophic bacteria) were obtained in the East China Sea in fall of 2000 and 2003. The average abundances of Synechococcus, Prochlorococcus, picoeukaryotes and heterotrophic bacteria were 10(5), 10(5), 10(4) and 10(6) cells ml(-1), respectively. Synechococcus, picoeukaryotes and heterotrophic bacteria were abundant at all the stations and presented higher concentration in the inner shelf where influences from the Changjiang effluent plumes and the coastal upwelling were evident, while Prochlorococcus was absent from the near-shore stations and became the dominant picophytoplankton population in offshore waters, where its abundance was comparable to that for heterotrophic bacteria. All picoplankton groups showed a reduction in cell number with depth, and a positive correlation with water temperature were observed, which reflected the importance of light and temperature on picoplankton growth. A negative relationship with salinity was found for heterotrophic bacteria along two sections across the East China Sea Shelf, and distribution of picoplankton was dominated by different water masses. The fixation could lead to loss in Prochlorococcus cell numbers within one month, and all the picoplankton numbers decreased dramatically after three months.     

Seasonality of top-down control of bacterioplankton at two central Red Sea sites with different trophic status

The role of bottom-up (nutrient availability) and top-down (grazers and viruses mortality) controls on tropical bacterioplankton have been rarely investigated simultaneously from a seasonal perspective. We have assessed them through monthly samplings over 2 years in inshore and offshore waters of the central Red Sea differing in trophic status. Flow cytometric analysis allowed us to distinguish five groups of heterotrophic bacteria based on physiological properties (nucleic acid content, membrane integrity and active respiration), three groups of cyanobacteria (two populations of Synechococcus and Prochlorococcus), heterotrophic nanoflagellates (HNFs) and three groups of viruses based on nucleic acid content. The dynamics of bacterioplankton and their top-down controls varied with season and location, being more pronounced in inshore waters. HNFs abundances showed a strong preference for larger prey inshore (r = −0.62 to −0.59, p = 0.001–0.002). Positive relationships between viruses and heterotrophic bacterioplankton abundances were more marked inshore (r = 0.67, p < 0.001) than offshore (r = 0.44, p = 0.03). The negative correlation between HNFs and viruses abundances (r = −0.47, p = 0.02) in shallow waters indicates a persistent seasonal switch between protistan grazing and viral lysis that maintains the low bacterioplankton stocks in the central Red Sea area.     

Plankton distribution across a slope current-induced front in the southern Bay of Biscay

Relatively warm (12.50–12.75°C) and high-salinity[<35.640 practical salinity units (PSU)] water flowing eastwardwas detected at the shelf-break during a cruise carried Outin the southern Bay of Biscay in Spring 1987. The slope currentinduced the formation of a convergent front separating well-mixedoceanic waters from haline-stratified coastal waters. Very highconcentrations of dissolved oxygen (295 µmol kg^–1)and chlorophyll a(>4.5 mg m^–3) were found at the outeredge of the frontal boundary. Small autotrophic flagellatesdominated the phytoplankton community. Primary production peakedat the boundary region. Estimated phytoplankton growth ratesindicated that active growth was taking place, with lower turnovertimes integrated over the water column at the frontal station(2.5–5 days) than at coastal (1.5–2.8 days) or oceanic(1.5–3.5 days) stations. The lowest doubling times (1–2days) were calculated for surface frontal populations. Accumulationof zooplankton was also observed associated with the convergentphysical structure, although this relationship was less markedthan for phytoplankton. Copepods, mainly Paracalanus parvus,Acartia clausi and Oithona helgolandica, formed the bulk ofthe mesozooplankton biomass. Compatibility between the sizeof phytoplankton cells and copepod size spectra indicate highfood availability for these animals, particularly in the vicinityof the front. The distribution of fish eggs and fish larvaewas also coupled with the slope current-induced front. Sardinelarvae were more abundant at the coastal side of the front,whereas larval stages of blue whiting reached the highest densitiesat off-shelf stations. Larvae of lamellibranch molluscs andbryozoa were restricted to nearshore waters, as the frontalboundary prevented larval dispersion to the open ocean. Theresults presented in this paper suggest that the Iberian slopecurrent and its associated shelf-break frontal structure werecrucial in controlling phytoplankton primary production, activityof grazers, distribution of larvae of fishes and benthic invertebrates,and ultimately in determining the structure of the pelagic foodweb in the southern Bay of Biscay during the seasonal periodof vertical mixing.     

Heterotrophic bacteria,zooflagellates, and ciliates in coastal waters of northeastern Black Sea

According to the parameters of density and biomass of heterotrophic bacterioplankton, waters in the ports of Novorossiysk and Tuapse were eutrophic-hypereutrophic and in the resort cities of Gelendzhik and Anapa they were hypereutrophic. The abundance of heterotrophic bacteria reached maximum values of 12.7–14.2 million cells/mL during the period of abnormally high water temperatures (August 2010) in recreational zones. Chainlike and filamentous forms (57–65%) of bacteria, which were typical for the strong coastal pollution of waters, prevailed. The abundance of zooflagellates (kinetoplastids) in semi-isolated water areas (port of Novorossiysk and Gelendzhik Bay) reached the level of highly eutrophic waters, 6.2–9.7 million ind/m³. However, the biomass of naked ciliates was 1.5 times lower than their maximum values earlier registered in coastal waters of the northeastern shelf. The peak of abundance of alien tintinnids of the genera Eutintinnus, Tintinnopsis, and Amphorellopsis, which were introduced via ballast waters of ships, was recorded in Novorossiysk Bay. The ratio of titinnids to the total abundance of ciliates increased 5 times and reached 25–40%. Protozoans developed poorly in the oil-polluted port of Tuapse and the open Anapa Bay.     

Presence of toxin-antitoxin systems in picocyanobacteria and their ecological implications

Picocyanobacteria (mainly Synechococcus and Prochlorococcus) contribute significantly to ocean’s primary production. Toxin-Antitoxin (TA) systems present in bacteria and archaea are known to regulate cell growth in response to environmental stresses. However, little is known about the presence of TA systems in picocyanobacteria. This study investigated complete genomes of Synechococcus and Prochlorococcus to understand the prevalence of TA systems in picocyanobacteria. Using the TAfinder software, Type II TA systems were predicted in 27 of 33 (81%) Synechococcus strains, but none of 38 Prochlorococcus strains contain TA genes. Synechococcus strains with larger genomes tend to contain more putative type II TA systems. The number of TA pairs varies from 0 to 42 in Synechococcus strains isolated from various environments. A linear correlation between the genome size and the number of putative TA systems in both coastal and freshwater Synechococcus was established. In general, open ocean Synechococcus contain no or few TA systems, while coastal and freshwater Synechococcus contain more TA systems. The type II TA systems inhibit microbial translation via ribonucleases and allow cells to enter the “dormant” stage in adverse environments. Inheritance of TA genes in freshwater and coastal Synechococcus could confer a recoverable persister mechanism important to survive in variable environments.Subject terms: Microbial ecology, Biooceanography     

Zooplankton diversity and the predatory impact by larval and small juvenile fish at the Fisher Banks in the North Sea

The biomass and diversity of the mesozooplankton and fish larvaecommunity were investigated across a frontal zone in the centralNorth Sea in the early summer, to investigate whether larvalfish predation is a regulator of mesozooplankton production.Pronounced changes in the mesozooplankton community were observedacross the front off the Jutlandic coast. The biomass and thediversity of copepods changed across the front as the populationof Calanus finmarchicus became abundant in the deeper water.The crustaceans (Acartia spp. and Evadne spp.) and polychaetelarvae dominated the mesozooplankton in the coastal water. Thebiomass of fish larvae was dominated by gadoid larvae. As inthe copepods, a shift in fish diversity was observed in thefrontal zone. On the coastal side of the front, whiting (Merlangiusmerlangus) dominated the biomass, while offshore from the frontwhiting were absent and cod (Gadus morhua) was the dominantlarval fish species on the deeper stations. The present investigationdemonstrates two different trophic pathways related to hydrographyin the central North Sea. First, in the shallow coastal water,the abundant small neritic copepods are predominately predatedby whiting, while in the offshore region the larger oceaniccopepods are predated by cod larvae. However, the predationpressure by the fish larvae was in general low (<10%) relativeto copepod production per day. Consequently, in the early summer,the copepod production potentially results in a build-up ofcopepod biomass on both sides of the front.     

Lake Superior Supports Novel Clusters of Cyanobacterial Picoplankton

Very little is known about the biodiversity of freshwater autotrophic picoplankton (APP) in the Laurentian Great Lakes, a system comprising 20% of the world's lacustrine freshwater. In this study, the genetic diversity of Lake Superior APP was examined by analyzing 16S rRNA gene and cpcBA PCR amplicons from water samples. By neighbor joining, the majority of 16S rRNA gene sequences clustered within the “picocyanobacterial clade” consisting of freshwater and marine Synechococcus and Prochlorococcus. Two new groups of Synechococcus spp., the pelagic Lake Superior clusters I and II, do not group with any of the known freshwater picocyanobacterial clusters and were the most abundant species (50 to 90% of the sequences) in samples collected from offshore Lake Superior stations. Conversely, at station Portage Deep (PD), located in a nearshore urbanized area, only 4% of the sequences belonged to these clusters and the remaining clones reflected the freshwater Synechococcus diversity described previously at sites throughout the world. Supporting the 16S rRNA gene data, the cpcBA library from nearshore station PD revealed a cosmopolitan diversity, whereas the majority of the cpcBA sequences (97.6%) from pelagic station CD1 fell within a unique Lake Superior cluster. Thus far, these picocyanobacteria have not been cultured, although their phylogenetic assignment suggests that they are phycoerythrin (PE) rich, consistent with the observation that PE-rich APP dominate Lake Superior picoplankton. Lastly, flow cytometry revealed that the summertime APP can exceed 10⁵ cells ml⁻¹ and suggests that the APP shifts from a community of PE and phycocyanin-rich picocyanobacteria and picoeukaryotes in winter to a PE-rich community in summer.     

Light-Stimulated Bacterial Production and Amino Acid Assimilation by Cyanobacteria and Other Microbes in the North Atlantic Ocean

We examined the contribution of photoheterotrophic microbes—those capable of light-mediated assimilation of organic compounds—to bacterial production and amino acid assimilation along a transect from Florida to Iceland from 28 May to 9 July 2005. Bacterial production (leucine incorporation at a 20 nM final concentration) was on average 30% higher in light than in dark-incubated samples, but the effect varied greatly (3% to 60%). To further characterize this light effect, we examined the abundance of potential photoheterotrophs and measured their contribution to bacterial production and amino acid assimilation (0.5 nM addition) using flow cytometry. Prochlorococcus and Synechococcus were abundant in surface waters where light-dependent leucine incorporation was observed, whereas aerobic anoxygenic phototrophic bacteria were abundant but did not correlate with the light effect. The per-cell assimilation rates of Prochlorococcus and Synechococcus were comparable to or higher than those of other prokaryotes, especially in the light. Picoeukaryotes also took up leucine (20 nM) and other amino acids (0.5 nM), but rates normalized to biovolume were much lower than those of prokaryotes. Prochlorococcus was responsible for 80% of light-stimulated bacterial production and amino acid assimilation in surface waters south of the Azores, while Synechococcus accounted for on average 12% of total assimilation. However, nearly 40% of the light-stimulated leucine assimilation was not accounted for by these groups, suggesting that assimilation by other microbes is also affected by light. Our results clarify the contribution of cyanobacteria to photoheterotrophy and highlight the potential role of other photoheterotrophs in biomass production and dissolved-organic-matter assimilation.     

Co-occurring Synechococcus ecotypes occupy four major oceanic regimes defined by temperature,macronutrients and iron

Marine picocyanobacteria, comprised of the genera Synechococcus and Prochlorococcus, are the most abundant and widespread primary producers in the ocean. More than 20 genetically distinct clades of marine Synechococcus have been identified, but their physiology and biogeography are not as thoroughly characterized as those of Prochlorococcus. Using clade-specific qPCR primers, we measured the abundance of 10 Synechococcus clades at 92 locations in surface waters of the Atlantic and Pacific Oceans. We found that Synechococcus partition the ocean into four distinct regimes distinguished by temperature, macronutrients and iron availability. Clades I and IV were prevalent in colder, mesotrophic waters; clades II, III and X dominated in the warm, oligotrophic open ocean; clades CRD1 and CRD2 were restricted to sites with low iron availability; and clades XV and XVI were only found in transitional waters at the edges of the other biomes. Overall, clade II was the most ubiquitous clade investigated and was the dominant clade in the largest biome, the oligotrophic open ocean. Co-occurring clades that occupy the same regime belong to distinct evolutionary lineages within Synechococcus, indicating that multiple ecotypes have evolved independently to occupy similar niches and represent examples of parallel evolution. We speculate that parallel evolution of ecotypes may be a common feature of diverse marine microbial communities that contributes to functional redundancy and the potential for resiliency.