Time series are critical to understand microbial plankton diversity and ecology

How diverse are marine planktonic protist communities? How much seasonality do they exhibit? For a very long time, these two old and challenging questions in the field of plankton ecology could be addressed only for large‐size protist species, based on cell counting under the microscope. The recent application of molecular techniques, notably massive marker‐gene amplicon sequencing approaches (metabarcoding), has allowed investigating with unprecedented level of resolution the small‐sized (<20 µm) planktonic eukaryotes too. An amazing diversity of these tiny organisms has been unveiled but details about their temporal dynamics remain much more elusive. In a From the Cover article in this issue of Molecular Ecology, Giner et al. (2019) introduce a new Recurrence Index (RI) to specifically look for seasonality in time‐series metabarcoding data. They inspected the temporal dynamics of all operational taxonomic units (OTUs) in a rich sequence data set of pico‐ and nanoplanktonic eukaryotes in samples collected monthly during 10 years. Although most OTUs did not show seasonality, some abundant ones did, which explains why some averaging methods can find seasonality at the less detailed level of whole planktonic communities. Not surprisingly, the very complex small‐sized eukaryotic plankton communities are composed of organisms with miscellaneous temporal dynamics.     

Does metazooplankton regulate the ciliate community in a shallow eutrophic lake?

1. As grazers on picoplankton and nanoplankton, planktonic ciliates form an important link in pelagic food webs. Ciliate communities may be controlled by predation by metazooplankton. In eutrophic systems, however, where the number of large crustaceans is often low, the mechanisms that regulate ciliate dynamics have rarely been described. 2. We conducted an enclosure experiment with natural and screened (145 μm) summer plankton communities to investigate the effect of the small‐sized crustacean zooplankton on ciliate community structure and the microbial loop in a shallow eutrophic lake. 3. The removal of the larger fraction of crustaceans initiated a decrease in total ciliate abundance. At the community level, we observed a substantial increase in large‐sized predacious ciliates (>100 μm) and a simultaneous decrease in the abundance of smaller ciliates (<20–40 μm) that were mostly bacterivores and bacterio‐herbivores. The compositional shift in the ciliate community, however, did not cascade down to the level of bacteria and edible phytoplankton.     

Spatial variability of marine bacterial and archaeal communities along the particulate matter continuum

Biotic and abiotic particles shape the microspatial architecture that defines the microbial aquatic habitat, being particles highly variable in size and quality along oceanic horizontal and vertical gradients. We analysed the prokaryotic (bacterial and archaeal) diversity and community composition present in six distinct particle size classes ranging from the pico‐ to the microscale (0.2 to 200 μm). Further, we studied their variations along oceanographic horizontal (from the coast to open oceanic waters) and vertical (from the ocean surface into the meso‐ and bathypelagic ocean) gradients. In general, prokaryotic community composition was more variable with depth than in the transition from the coast to the open ocean. Comparing the six size‐fractions, distinct prokaryotic communities were detected in each size‐fraction, and whereas bacteria were more diverse in the larger size‐fractions, archaea were more diverse in the smaller size‐fractions. Comparison of prokaryotic community composition among particle size‐fractions showed that most, but not all, taxonomic groups have a preference for a certain size‐fraction sustained with depth. Species sorting, or the presence of diverse ecotypes with distinct size‐fraction preferences, may explain why this trend is not conserved in all taxa.     

Unexpected biodiversity of ciliates in marine samples from below the photic zone

Marine microbial eukaryotes play critical roles in planktonic food webs and have been described as most diverse in the photic zone where productivity is high. We used high‐throughput sequencing (HTS) to analyse the spatial distribution of planktonic ciliate diversity from shallow waters (<30 m depth) to beyond the continental shelf (>800 m depth) along a 163 km transect off the coast of New England, USA. We focus on ciliates in the subclasses Oligotrichia and Choreotrichia (class Spirotrichea), as these taxa are major components of marine food webs. We did not observe the decrease of diversity below the photic zone expected based on productivity and previous analyses. Instead, we saw an increase of diversity with depth. We also observed that the ciliate communities assessed by HTS cluster by depth layer and degree of water column stratification, suggesting that community assembly is driven by environmental factors. Across our samples, abundant OTUs tend to match previously characterized morphospecies while rare OTUs are more often undescribed, consistent with the idea that species in the rare biosphere remain to be characterized by microscopy. Finally, samples taken below the photic zone also reveal the prevalence of two uncharacterized (i.e. lacking sequenced morphospecies) clades – clusters X₁ and X₂ – that are enriched within the nano‐sized fraction (2–10 μm) and are defined by deletions within the region of the SSU‐rDNA analysed here. Together, these data reinforce that we still have much to learn about microbial diversity in marine ecosystems, especially in deep‐waters that may be a reservoir for rare species and uncharacterized taxa.     

Major imprint of surface plankton on deep ocean prokaryotic structure and activity

Deep ocean microbial communities rely on the organic carbon produced in the sunlit ocean, yet it remains unknown whether surface processes determine the assembly and function of bathypelagic prokaryotes to a larger extent than deep‐sea physicochemical conditions. Here, we explored whether variations in surface phytoplankton assemblages across Atlantic, Pacific and Indian ocean stations can explain structural changes in bathypelagic (ca. 4,000 m) free‐living and particle‐attached prokaryotic communities (characterized through 16S rRNA gene sequencing), as well as changes in prokaryotic activity and dissolved organic matter (DOM) quality. We show that the spatial structuring of prokaryotic communities in the bathypelagic strongly followed variations in the abundances of surface dinoflagellates and ciliates, as well as gradients in surface primary productivity, but were less influenced by bathypelagic physicochemical conditions. Amino acid‐like DOM components in the bathypelagic reflected variations of those components in surface waters, and seemed to control bathypelagic prokaryotic activity. The imprint of surface conditions was more evident in bathypelagic than in shallower mesopelagic (200–1,000 m) communities, suggesting a direct connectivity through fast‐sinking particles that escape mesopelagic transformations. Finally, we identified a pool of endemic deep‐sea prokaryotic taxa (including potentially chemoautotrophic groups) that appear less connected to surface processes than those bathypelagic taxa with a widespread vertical distribution. Our results suggest that surface planktonic communities shape the spatial structure of the bathypelagic microbiome to a larger extent than the local physicochemical environment, likely through determining the nature of the sinking particles and the associated prokaryotes reaching bathypelagic waters.     

Contribution of ciliated protozoa to the planktonic biomass in a series of Ontario lakes: quantitative estimates and dynamical relationships

The plankton of nine Ontario lakes spanning several physiographicregions was sampled every two weeks during the ice-free periodof 1981, and one lake was studied in the three previous years.Phytoplankton, zooplankton, and ciliated protozoa were sampled,counted and sized. The size data were converted to biomass estimatesto yield quantitative comparisons of the relative allocationof biomass among different functional compartments. This isthe first study to look simultaneously and quantitatively atthe total plankton system of lakes (including ciliates, pbytoplanktonand net zooplankton) over a broad physiographic region. Ciliatesconstitute –10% of the non-algal biomass and 5% of thetotal planktonic biomass of these lakes. Ciliate standing cropsamong lakes are significantly corrrelated with total organicand total inorganic carbon concentrations in the water column,while the dynamics of ciliate biomass fluctuations are significantlycorrelated with variations in total phosphorus concentration,in conductivity, in Kjeldahl nitrogen concentration, and ininorganic carbon content. There appears to be a significantdynamical relationship between ciliates as a proportion of thetotal planktonic biomass, exclusive of filamentous and large(>30 µm) spherical algae, and the relative biomassof small algae (2–5 µm) as a fraction of total algalbiomass, again exclusive of filaments and large (>30 µm)algae. The hypothesis is advanced that ciliates primarily functionas bacterial grazers in planktonic ecosystems and that theirprimary competitors in this role are rotifers.     

High abundance of picoplankton-ingesting ciliates during late fall in Lake Kinneret, Israel

Small, aloricate ciliates dominated the biomass of heterotrophicprotists throughout the water column at the end of the periodof stratification in Lake Kinneret, Israel The integrated biomassof cilates was 5–20 times that of heterotrophic flagellatesDuring incubation experiments, ciliate growth rates in cpilimneticwater corresponded to population doubling times of 9.6–19.4h, while flagellate populations showed no growth. Most of thealiates were small forms (10–30 µm long), includingscuticocihates, choreotnchs, Coleps spp. and Colpoda spp., andappeared to be consuming bacteria, coccoid cyanobacteria, and<5 µm eukaryotic algae. Grazing rates of cihate assemblageson picoplankton in the epilimnion, as determined by the uptakeof fluorescently labeled bacteria and cyanobactena, ranged from62 to 86 nl cell^–1 h^–1 Colpoda steini, isolatedfrom lakewater, grew on a cultured freshwater Synechococcussp with a doubling time of 4.5 h, and a gross growth efficiencyof 48% The estimated daily requirements of ciliates for growthapproximately equalled total phytoplankton production. We calculatedthat ciliates in the epilimnion were clearing 4–10% ofthe bacterioplankton and cyanobactenal standing stocks per daySince this would not be sufficient food consumption to meetdaily carbon requirements of the aliates, it is likely thatthese organisms were also grazing a significant amount of autotrophicand heterotrophic eukaryotic cells in Lake Kinneret.     

Past President's Address: Protists of the Mesopelagic and a Bit on the Long Path to the Deep Sea

The deep sea has long been a mysterious and attractive habitat for protistologists. However, logistical difficulties severely limit sampling opportunities. Consequently, our knowledge of the protists in the deep sea, (arguably the largest habitat on earth), is relatively sparse. Here, we present a unique time‐series concerning three different protist taxa that share only the characteristics of being relatively large, robust to sampling, and easily identifiable to species level using light microscopy: tintinnid ciliates, phaeogromid cercozoans (e.g. Challengerids) and amphisolenid dinoflagellates. We sampled a near‐shore deep water site in the N.W. Mediterranean Sea at 250 m depth over a 2‐yr period at approximately weekly intervals from January 2017 to December 2018. To our knowledge, no previous studies have employed sampling on a similar time scale. We found taxa that appear to be restricted to deep waters, distinct seasonal patterns of abundance in some taxa, and in others nonseasonal successional patterns. Based on data from sampling following a flash flood event, the Challengerid population appeared to respond positively to a pulse of terrigenous input. Some of the distinct mesopelagic tintinnid ciliates and amphisolinid dinoflagellates were also found in two samples from the North Atlantic mesopelagic gathered from near the Azores Islands in September 2018. We conclude that there are a variety of protist taxa endemic to the mesopelagic, that the populations are dynamic, and they may be widely distributed in the deep waters of the world ocean.     

Quantifying long‐term recurrence in planktonic microbial eukaryotes

How much temporal recurrence is present in microbial assemblages is still an unanswered ecological question. Even though marked seasonal changes have been reported for whole microbial communities, less is known on the dynamics and seasonality of individual taxa. Here, we aim at understanding microbial recurrence at three different levels: community, taxonomic group and operational taxonomic units (OTUs). For that, we focused on a model microbial eukaryotic community populating a long‐term marine microbial observatory using 18S rRNA gene data from two organismal size fractions: the picoplankton (0.2–3 µm) and the nanoplankton (3–20 µm). We have developed an index to quantify recurrence in particular taxa. We found that community structure oscillated systematically between two main configurations corresponding to winter and summer over the 10 years studied. A few taxonomic groups such as Mamiellophyceae or MALV‐III presented clear recurrence (i.e., seasonality), whereas 13%–19% of the OTUs in both size fractions, accounting for ~40% of the relative abundance, featured recurrent dynamics. Altogether, our work links long‐term whole community dynamics with that of individual OTUs and taxonomic groups, indicating that recurrent and non‐recurrent changes characterize the dynamics of microbial assemblages.     

Oldest fossil ciliates from the Cryogenian glacial interlude reinterpreted as possible red algal spores

The Cryogenian Period experienced two long lived global glaciations known as Snowball Earths. While these events were dramatic, eukaryotic life persisted through them, and fossil evidence shows that eukaryotes thrived during the c. 30-million-year interlude between the glaciations. Carbonate successions have become an important taphonomic window for this interval. One of the most notable examples is the c. 662–635 Ma Taishir Formation (Tsagaan Olom Group, Zavkhan Terrane, Mongolia) which has yielded a number of eukaryotic fossil taxa. Here, we examine more closely the morphology and taxonomic affinity of some of these Taishir fossils previously interpreted as remains of ciliate tintinnid loricae (purportedly the oldest fossil ciliates). New morphological and ultrastructural analyses indicate that these fossils are not ciliate tintinnids. Instead, we propose a new interpretation: that they are algal reproductive structures related to coeval macroscopic organic warty sheets described as putative red algae. We report the first occurrence of these fossils in the earliest Ediacaran Ol Formation, indicating that this taxon persisted through the Marinoan Snowball Earth. A new interpretation of these fossils as putative red algal spores has broad implications for our understanding of biodiversity in the Neoproterozoic Era, specifically during the Cryogenian Period, and for the antiquity of ciliates.     

Pelagic ciliated protozoa in two monomictic, southern temperate lakes of contrasting trophic state: seasonal distribution and abundance

Two lakes of contrasting trophic state in the central NorthIsland of New Zealand were sampled monthly for protozoan ciliatesand potential food resources. Oligotrichs dominated numbersin both lakes. Subdominants in oligotrophic Lake Taupo includedAskenasia, Pscudobalanion and Urotri-cha, and in eutrophic LakeOkaro Prorodon, Coleps, Urocentrum, Stentor and Spirostomumwere important. Biomass was dominated by large predatory ciliatesand Stentor in Lake Taupo, and Prorodon and Stentor in LakeOkaro. The importance of Prorodon and Stentor to ciliate biomassis unusual and has not been reported for northern hemispherelakes. Small ciliates (<20 µm) capable of consumingparticles <2 µm were a major component of the ciliatecommunity in Lake Taupo. Peaks in ciliate abundance occurredat the same time in both lakes: in autumn, at the beginningof mixis and in spring. Ciliates were vertically stratifiedduring mixis and stratification in both lakes. The effect wasmore pronounced during deoxygenation of the hypolimnion in LakeOkaro which excluded oligotrichs and introduced benthic ciliates.Ciliates were less abundant (mean 40001^–1 in Lake Okaroand 9001^–1 in Lake Taupo) than in comparable northerntemperate lakes. There was no correlation between the seasonaldistribution of ciliates and chlorophyll a, primarily causedby a winter peak in chlorophyll a dominated by large speciesof phytoplankton in Lake Taupo, at a time when ciliate numberswere low. The only consistent, significant correlations weretotal ciliate numbers and individual species of ciliates withbacterial concentrations in both lakes and with picophytoplanktonin Lake Taupo.     

Microbial eukaryote life in the new hypersaline deep-sea basin Thetis

Only recently, a novel anoxic hypersaline (thalassic) basin in the eastern Mediterranean was discovered at a depth of 3,258 m. The halite-saturated brine of this polyextreme basin revealed one of the highest salt concentrations ever reported for such an environment (salinity of 348‰). Using a eukaryote-specific probe and fluorescence in situ hybridization, we counted 0.6 × 10⁴ protists per liter of anoxic brine. SSU rRNA sequence analyses, based on amplification of environmental cDNA identified fungi as the most diverse taxonomic group of eukaryotes in the brine, making deep-sea brines sources of unknown fungal diversity and hotspots for the discovery of novel metabolic pathways and for secondary metabolites. The second most diverse phylotypes are ciliates and stramenopiles (each 20%). The occurrence of closely related ciliate sequences exclusively in other Mediterranean brine basins suggests specific adaptations of the respective organisms to such habitats. Betadiversity-analyses confirm that microeukaryote communities in the brine and the interface are notably different. Several distinct morphotypes in brine samples suggest that the rRNA sequences detected in Thetis brine can be linked to indigenous polyextremophile protists. This contradicts previous assumptions that such extremely high salt concentrations are anathema to eukaryotic life. The upper salinity limits for eukaryotic life remain unidentified.     

Climate change-predicted shifts in the temperature regime of shallow groundwater produce rapid responses in ciliate communities

The ciliates living in a shallow groundwater system in southern Ontario, Canada were subjected to an in situ temperature manipulation over 14 months. Ciliates were collected from the bed surface of a small springbrook and from interstitial water collected at five depths beneath its surface. Mean temperature elevations established at each depth (?20, ?40, ?60, ?80, and ?100 cm) between the experiment's control and treatment blocks were 1.9, 3.5, 3.9, 3.8, and 3.6 °C, respectively, and were based on global warming projections for the region. In total, 160 species of ciliate belonging to 85 genera were identified. Overall, the treatment block had a higher density (6510±342 cells L^?1; ±1 SE) than the control (5797±237 cells L^?1), but densities were both vertically and longitudinally variable. Control densities decreased with depth, whereas treatment densities were more equal among depths. Total species richness showed no significant difference between blocks when combining all sampling dates and depths, although species composition changed. The ciliate community was dominated by small (15–50 μm), followed by medium (50–200 μm), and only a few large‐sized (>200 μm) species. Small ciliates contributed 82–97% of the total density. Small ciliates also contributed more to the treatment (94%) than the control block (88%). The most common ciliate feeding groups were bacterivores, omnivores, predators, and algae‐diatom feeders, with bacterivores being most dominant (83–99% of the total numbers collected). Ordination analyses revealed that ciliate distribution was strongly correlated with groundwater temperature, although dissolved oxygen level, concentrations of ammonia and nitrate, and depth also appeared to be influential. Peak densities of many species occurred in either the control or treatment blocks, but not in both. The benefits of using ciliates as a proxy for higher, much longer‐lived, eukaryotes in climate change studies are discussed.     

Effects of Daphnia on microzooplankton communities

Intact phytoplankton and microzooplankton communities from eutrophicStar Lake were incubated for 4 days with and without Daphniapulex, Daphnia galeaia mendotae, or a natural assemblage ofDaphnia species. They were sampled at the onset and terminationof the experiment for bacterial, phytoplankton, ciliate, rotifer,copepod and cladoceran densities. The cladocerans had variedeffects on the rotifers, ranging from significant suppressionof most rotifer species (Keratella cochlearis, Polyarthra remata,Keratella crassa) in the D.pulex jars, to the suppression ofone (K.crassa) or no species in the D.galeata mendotae and StarLake Daphnia assemblage jars, respectively. Small ciliates (<30µm, longest dimension), such as Strobilidium sp. and Pseudo-cyclidiumsp., were adversely affected by most of the cladoceran treatments,while several larger ciliates (>81 µm) were unaffectedin all such treatments. Ciliates were not consistently morevulnerable to cladoceran suppression than similarly sized rotifers.The suppression of ciliates and rotifers was attributable toboth direct effects (predation, interference, or both) and indirecteffects (e.g. resource competition) of the cladocerans. ¹Present address: Department of Biology, University of Louisville,Louisville, KY 40292, USA     

The Size of DNA Molecules and Chromatin Organization in the Macronucleus of the Ciliate Didinium nasutum (Ciliophora)

Pulsed‐field gel electrophoresis (PFGE) was applied to analyze the molecular karyotype of the ciliate Didinium nasutum. The data obtained indicate that D. nasutum belongs to the ciliate species with subchromosomal macronuclear genome organization. No short “gene‐sized” DNA molecules were detected. Macronuclear DNAs formed a continuous spectrum from 50 kbp to approximately 1,000 kbp in size with a peak plateau between 250 and 400 kbp. The macronuclear DNA molecules were packed into chromatin bodies of 80–265 nm in size. Comparison of the PFGE and electron microscopic data shows that most if not all chromatin bodies contain more than one DNA molecule.     

A mechanism of transmission and factors affecting coral susceptibility to <Emphasis Type="Italic">Halofolliculina</Emphasis> sp. infection

Anecdotal evidence collected since 2004 suggests that infections caused by ciliates in the genus Halofolliculina may be related to coral mortality in more than 25 scleractinian species in the Caribbean. However, the relationship between the presence of ciliates and coral mortality has not yet been firmly established. Field and laboratory manipulations were used to test if ciliate infections harm corals, if ciliates are able to infect healthy colonies, and if coral susceptibility to ciliate infection depends on temperature, depth, distance to an infected colony, and the presence of injuries. Ciliate infections were always characterized by a visually detectable front of ciliates located on recently exposed coral skeletons. These infections altered the normal structure of the colony by causing tissue mortality (0.8 ± 0.95 cm month⁻¹, mean ± SD) and by delaying or preventing recovery from injuries. Under laboratory conditions, ciliates transmitted directly and horizontally from infected to healthy hosts, and coral susceptibility to ciliate infections increased with the presence of injuries. After invasion, the ciliate population grew, rapidly and after 8 d, produced tissue mortality on 32% of newly infected hosts. Thus, our results support the existence of a new Caribbean coral syndrome that is associated with tissue mortality, is infectious, and transmits directly and horizontally. Even though the role of ciliates in the development of lesions on coral tissues remains unclear, their presence is by far the most conspicuous sign of this syndrome; thus, we propose to name this condition Caribbean ciliate infection (CCI). Communicated by Biology Editor Dr Michael Lesser     

Annual Changes of Abundance and Biomass of Planktonic Ciliates in the Gdańsk Basin,Southern Baltic

Seasonal changes in the species composition, abundance and biomass of planktonic ciliates were determined every 2–3 weeks at two sites of 30 m depth and one location of 105 m depth in the southwestern Gdańsk Basin between January 1987 and January 1988. A total of 40 ciliate taxa were observed during this period. Autotrophic Mesodinium rubrum dominated ciliate abundance and biomass: maximal values of 50 · 10⁻¹ ind. ^1-1 and 65 μg C 1⁻¹ were recorded. The annual mean biomass of M. rubrum comprised 6 to 9% of the annual mean phytoplankton biomass. The highest abundances and biomasses of heterotrophic ciliates were noted at all stations in the spring and summer in the euphotic zone with maximum values of 28 · 10³ ind. 1⁻¹ and 23 μg C 1⁻¹. Three ciliates assemblages were distinguished in the epipelagic layer: large and medium-size non-predatory ciliates, achieving peak abundance in spring and autumn; small-size microphagous ciliates and epibiotic ciliates which were abundant in summer, and large-size predacious ciliates dominating in spring. Below 60 m, a separate deep-water ciliate community composed of Prorodon-like ciliates and Metacystis spp. was found. The ciliate biomass in the 60–105 m layer was similar to the ciliate biomass in the euphotic zone. The heterotrophic ciliate community contributed 10 to 13% to the annual mean zooplankton biomass. The potential annual production of M. rubrum comprised 6 to 9% of the total primary production. Carbon demand of non-predatory ciliates, calculated on the basis of their potential production, was estimated to be equivalent to 12–15% of the gross primary production.     

Stratification and dynamics of microbial loop communities in Lake Fryxell, Antarctica

1. Lake Fryxell, situated in the McMurdo Dry Valleys, Antarctica, offers the opportunity to study microbial loop processes in the absence of crustacean zooplankton and other higher organisms. This is the first study of Lake Fryxell to provide detailed temporal and vertical variations of microbial loop organisms.
2. Protozoan communities are concentrated around the chemocline (9–10 m) in Lake Fryxell. Phototrophic nanoflagellates (PNAN), heterotrophic nanoflagellates (HNAN) and ciliates formed deep maxima of 14 580, 694 and 58 cells mL−1 respectively. Although abundance and biomass at the chemocline was high, diversity of protozoa was low, Plagiocampa accounting for> 80% of the total ciliate biomass.
3. In the mixolimnion (4.5–8 m), protozoa were less abundant, but more diverse, with 24 ciliate morphotypes being identified within this region of the water column. Inter-annual variability of protozoan biomass and abundance was greater in the mixolimnion than at the chemocline due to more variable nutrient and prey concentrations.
4. Physicochemical gradients in Lake Fryxell were very stable because the perennial ice cover reduced wind driven currents. As a consequence, ciliate species occurred in distinct depth strata, Monodinium being most abundant directly beneath the ice cover, Askenasia having maximum abundance at 8 m and Plagiocampa dominating ciliate biomass at the chemocline. The lack of vertical mixing reduced seasonal successions of PNAN and ciliate species. Three cryptophyte species dominated the PNAN community at all times (>79% of total biomass).     

Feeding by larval and post-larval ctenophores on microzooplankton

Feeding by the coastal ctenophorc, Mnemiopsis leidyi, on microplanktonwas investigated. Larval ctenophores (tentaculate stage) grewbest and had the highest survival rates when offered a mixtureof ciliates and copepod nauplii. Larvae did not survive whenoffered phytoplankton alone. Clearing of planktonjc ciliatesby post-larval ctenophores was a function of the ciliate speciesand the size of the predator. Removal of small ciliates (<20µm in size) and phytoplankton was negligible. Small post-larvalctenophores (volume <4 cm³) had higher biovolume-specificclearing rates (0.5–1.5 1 cm^–3 day^–1) thandid larger ctenophores fed the same ciliate species. Duringin situ incubations, adult M. leidyi removed ciliates, rotifersand copepod nauplii from natural microplankton assemblages.The data indicate that non-crustacean microzooplanlctoo arean important component of the diet of larval and post-larvallocate cteoophores, particularly when copepod standing stocksare low.     

Prevalent Ciliate Symbiosis on Copepods: High Genetic Diversity and Wide Distribution Detected Using Small Subunit Ribosomal RNA Gene

Toward understanding the genetic diversity and distribution of copepod-associated symbiotic ciliates and the evolutionary relationships with their hosts in the marine environment, we developed a small subunit ribosomal RNA gene (18S rDNA)-based molecular method and investigated the genetic diversity and genotype distribution of the symbiotic ciliates on copepods. Of the 10 copepod species representing six families collected from six locations of Pacific and Atlantic Oceans, 9 were found to harbor ciliate symbionts. Phylogenetic analysis of the 391 ciliate 18S rDNA sequences obtained revealed seven groups (ribogroups), six (containing 99% of all the sequences) belonging to subclass Apostomatida, the other clustered with peritrich ciliate Vorticella gracilis. Among the Apostomatida groups, Group III were essentially identical to Vampyrophrya pelagica, and the other five groups represented the undocumented ciliates that were close to Vampyrophrya/Gymnodinioides/Hyalophysa. Group VI ciliates were found in all copepod species but one (Calanus sinicus), and were most abundant among all ciliate sequences obtained, indicating that they are the dominant symbiotic ciliates universally associated with copepods. In contrast, some ciliate sequences were found only in some of the copepods examined, suggesting the host selectivity and geographic differentiation of ciliates, which requires further verification by more extensive sampling. Our results reveal the wide occurrence and high genetic diversity of symbiotic ciliates on marine copepods and highlight the need to systematically investigate the host- and geography-based genetic differentiation and ecological roles of these ciliates globally.