Community structure, biomass and productivity of size-fractionated summer phytoplankton populations in lower Narragansett Bay, Rhode Island

Seventeen size-fractionation experiments were carried out duringthe summer of 1979 to compare biomass and productivity in the< 10, <8 and <5 µm size fractions with that ofthe total phytoplankton community in surface waters of NarragansettBay. Flagellates and non-motile ultra-plankton passing 8 µmpolycarbonate filters dominated early summer phytoplankton populations,while diatoms and dinoflagellates retained by 10 µm nylonnetting dominated during the late summer. A significant numberof small diatoms and dinoflagellates were found in the 10–8µm size fraction. The > 10 µm size fraction accountedfor 50% of the chlorophyll a standing crop and 38% of surfaceproduction. The <8 µm fraction accounted for 39 and18% of the surface biomass and production. Production by the< 8 µm fraction exceeded half of the total communityproduction only during a mid-summer bloom of microflagellates.Mean assimilation numbers and calculated carbon doubling ratesin the <8 µm (2.8 g C g Chl a^–1 h^–1; 0.9day^–1)and<5 µm(1.7 g C g Chl a^–1h^–1; 0.5day^–1)size fractions were consistently lower than those of the totalpopulation (4.8 g C g Chl a^–1 h^–1; 1.3 day^–1)and the <10 µm size fraction (5.8 g C g Chl a^–1h^–1; 1.4 day ^–1). The results indicate that smalldiatoms and dinoflagellates in fractionated phytoplankton populationscan influence productivity out of proportion to their numbersor biomass. ¹Present address: Australian Institute of Marine Science, P.M.B.No. 3, Townsville M.S.O., Qld. 4810, Australia.     

Ctenophore-zooplankton-phytoplankton interactions in Narragansett Bay, Rhode Island, USA, during 1972-1977

Plankton dynamics at a station in lower Narragansett Bay, RIare compared for six summer and fall seasons, 1972–1977.In four of these years, initiation of the summer pulse of thectenophore Mnemiopsis leidyi was accompanied by a rapid declinein zooplankton abundance and a summer phytoplankton bloom. Terminationof the phytoplankton bloom coincided with depleted ctenophoreabundance and increased zooplankton biomass in two of the years.Yearly variations in the summer abundance of the diatom Skeletonemacostatum were positively related to the magnitude of the ctenophorepulse. The magnitude of ctenophore population was related tothe zooplankton biomass present at the start of the pulse. Theserelationships, the timing and magnitude of the plankton eventssuggest that M. leidyi regulated summer zooplankton and phytoplanktondynamics. Ctenophores may control phytoplankton blooms indirectlythrough their predation on herbivorous zooplankton and directlyby the nutrient excretion accompanying such grazing. This evidencethat a planktonic carnivore two trophic steps removed from thephytoplankton regulates the latter's dynamics in NarragansettBay is analogous to reported regulation of benthic algal (kelp)dynamics by the sea otter, lobster and various crabs throughtheir predation on herbivorous sea urchins. The factors responsiblefor the seasonal decrease in ctenophores remain unresolved;ctenophore predators on Mnemiopsis are absent in NarragansettBay. Infection by the vermiform larval anemone, Edwardsia lineata,grazing by the butterfish, Peprilus triacanthus, and changesin food availability, temperature and salinity likewise do notexplain this disappearance.     

Seasonal variations of tintinnids (Ciliophora: Oligotrichida) in Narragansett Bay, Rhode Island, U.S.A.

Tintinnids exhibit a bimodal peak of abundance in NarragansettBay, with a minimum in late spring and a lesser peak in latesummer-early autumn, depending on location. Thirty-three speciesin eight genera were identified, with the fauna dominated bythe genus Tintinnopsis. Tintinnid abundance was not obviouslycorrelated with salinity, temperature, or nanoplankton chlorophyllconcentration. Patterns of change in seasonal abundance canserve as a framework for posing additional questions of therole tintinnids play in marine food webs.     

Nitrogen dynamics in lower Narragansett Bay. II. Phytoplankton uptake, depletion rates of nitrogenous nutrient pools, and estimates of ecosystem remineralization

Phytoplankton nitrogen demand in lower Narragansett Bay, RhodeIsland, measured during the winter-spring of 1977–78 andsummers of 1978 and 1979, is compared with estimates of zooplanktonand the benthic nitrogen remineralization drawn from the resultsof ekperimental field studies. Measured uptake rates would generallylead to the depletion of available nitrogenous nutrient stockswithin hours, and usually exceeded estimates of benthic pluszooplankton remineralization. Additional estimates of nitrogeninputs from sewage and riverine sources appear insufficientto make up the difference. The discrepancy lends support tothe paradigm that water column remineralization by microheterotrophsmay supply much, if not most, of the nitrogen needs of coastalphytoplankton.     

Ploidy Distribution of the Harmful Bloom Forming Macroalgae Ulva spp. in Narragansett Bay,Rhode Island,USA, Using Flow Cytometry Methods

Macroalgal blooms occur worldwide and have the potential to cause severe ecological and economic damage. Narragansett Bay, RI is a eutrophic system that experiences summer macroalgal blooms composed mostly of Ulva compressa and Ulva rigida, which have biphasic life cycles with separate haploid and diploid phases. In this study, we used flow cytometry to assess ploidy levels of U. compressa and U. rigida populations from five sites in Narragansett Bay, RI, USA, to assess the relative contribution of both phases to bloom formation. Both haploid gametophytes and diploid sporophytes were present for both species. Sites ranged from a relative overabundance of gametophytes to a relative overabundance of sporophytes, compared to the null model prediction of √2 gametophytes: 1 sporophyte. We found significant differences in cell area between ploidy levels for each species, with sporophyte cells significantly larger than gametophyte cells in U. compressa and U. rigida. We found no differences in relative growth rate between ploidy levels for each species. Our results indicate the presence of both phases of each of the two dominant bloom forming species throughout the bloom season, and represent one of the first studies of in situ Ulva life cycle dynamics.     

Experimental evaluation of herbivory in the ctenophore Mnemiopsis leidyi relevant to ctenophore-zooplankton-phytoplankton interactions in Narragansett Bay, Rhode Island, USA

Herbivory of Mnemiopsis leidyi and its interactions with phytoplanktonand non-gelatinous zooplankton were examined in small-scalemicrocosm experiments. Clearance rates for M. leidyi incubatedwith phytoplankton were generally negative, but ranged up to4.5 1 ctenophore^–1 day^–1 when the large (80 µmø) diatom Ditylum brightwelli was offered as a food source.These highest ingestion rates would provide Mnemiopsis withonly 21 % of its daily carbon requirements for respiration.Mean shrinkage of M. leidyi was 8.2–51% when incubatedwith phytoplankton. Although M. leidyi neither fed activelyon phytoplankton, nor satisfied its nutritional needs on sucha diet, the chain-forming diatom Skeletonema costatum becameentangled in mucus strands and balls produced by M. leidyi inthe absence of zooplankton. Attachment onto mucus occurred atphytoplankton concentrations commonly observed in NarragansettBay and may be important in the formation of "marine snow" duringsummer M. leidyi pulses; phytoplankton sinking rate and the"package size" available to herbivores would also be affected.The experiments support our previous hypothesis based on fieldobservations in Narragansett Bay that M. leidyi indirectly regulatesphytoplankton abundance there during the summer bloom as a consequenceof predation on zooplankton. The extent to which M. leidyi influencedphytoplankton dynamics in the microcosms was dependent on therelative abundance and physiological state of the three trophiclevels. A food web diagram for M. leidyi is presented.     

The contribution of ammonia excreted by zooplankton to phytoplankton production in Narragansett Bay

Ammonia excreted by mixed zooplankton populations over an annual(1972–1973) cycle in Narragansett Bay varied from 0.04to 3.21 µg at NH₃-N dry wt^–1 day^–1, exclusiveof two exceptional rates measured one year apart: 11.74 and18.39 µg at NH³-N mg dry wt^–1 day^–1. Grossphytoplankton production integrated over the year (1972–1973)averaged 151 mg C m^–3 day^–1 for an 8 m water column;peaks of 332 and 905 mg C m^–3 day^–1 occurred duringthe winter-spring and summer blooms, respectively. Excretedammonia, integrated seasonally and annually, contributed only0.2% and 4.9% of the nitrogen required for observed gross productionduring the winter-spring and summer blooms, respectively, and4.4% annually. However, excreted ammonia may be an importantsource of the nitrogen required by Skeletonema costatum, thedominant diatom in Narragansett Bay, during the post-bloom periodwhen 186% of the nitrogen required for its net production wasmet by ammonia excretion. A combination of zooplankton ammoniaexcretion and benthic ammonia flux contributed 22% of the nitrogenrequired for the annual gross production (440 g C m^–2)while 51% of the nitrogen required for the net production ofSkeletonema was accounted for by regenerated nitrogen. ¹This research was supported by NSF grant GA 31319X awardedto Dr.T.J.Smayda.     

Monterey Bay phytoplankton I. Seasonal cycles of phytoplankton assemblages

Netplankton collections from shallow-water stations in MontereyBay, California were examined for 1976 and 1977. The predominatingphytoplankton forms were neritic diatom species. Recurrent speciesgroups were identified using Fager's regroup analysis and theirdistributions related to hydrographic seasons. Neritic groupswere relatively more abundant during the Davidson Current andupweiling periods; however, many of the species were presentthroughout much of the year. The appearance of oceanic groupsand species was seasonal and associated with advective events. One neritic species group was of overwhelming numerical importancebut within the group conspicuous changes in relative abundanceof phytoplankton populations occurred seasonally among speciesfrom this group. This group appears to be composed of residentspecies and may be locally retained as vegetative cells or benthicresting stages. Species cycles in Monterey Bay may be regardedas largely the result of suc-cessional changes or cycles ofautochthonous species, while species introduced by adjectiveprocesses appear to be of minor importance. The study has suggested that the presence of species in a ‘seedpopulation’, either as vegetative cells or benthic restingstages, may be important as the growth rate of the species inunderstanding and predicting composition and relative abundancein coastal phytoplankton assemblages.     

Toxic and noxious phytoplankton in Big Glory Bay,Stewart Island,New Zealand

Diurnal vertical profile sampling of the water column, during a fish killing bloom of the raphidophycean alga Heterosigma akashiwo, revealed a phytoplankton population otherwise composed almost entirely of a variety of dinoflagellates. Of these Glenodinium danicum, Dinophysis acuta, Polykrikos schwartzii, Ceratium furca and Gyrodinium spirale were predominant. The distribution of the major species within the phytoplankton were documented and evidence of synchronous vertical migration of H. akashiwo, G. danicum and P. schwartzii was observed. Extracts of shellfish obtained during the bloom and tested by mouse bioassay showed no PSP toxicity but a marginal degree of DSP toxicity. During a subsequent one year phytoplankton monitoring programme another potentially noxious species (Chaetoceros convolutus) appeared and the seasonal reoccurrence of species present during the bloom (e.g. H. akashiwo) was observed. Important year to year differences in the summer phytoplankton (diatom versus flagellate dominated populations) were apparent and analysis of climate data showed that these differences related to different weather conditions prevailing during the two summer periods sampled. The data suggest the fish killing bloom was giving a chance to develop by a prolonged period of warm, calm weather (during which several heavy rainfall events occurred) leading to stable hydrographic conditions (i.e. stratification) and an increase in the retention time of water within the bay.     

Temporal patterns and variations in phytoplankton community organization and abundance in Narragansett Bay during 1959-1980

Incident irradiance, surface water temperature and phytoplanktonspecies abundances were measured at weekly intervals in NarragansettBay from 1959 through 1980. Stepwise discriminant analyses (SDA)of this 22-year data set indicate that fundamental ecosystemchanges occurred between the 1960s and 1970s, with 1969 beingthe key transitional year in these decadal shifts in phyto planktontaxonomic structure and seasonal abundance. This decadal shiftwas accompanied by the increased summer abundance of small Thalassiosiraspp., which first appeared in 1966 and by 1969 became the sixthmost important phytoplankton component in this bay. Decadaltrends in phyto plankton community organization and abundancewere also accompanied by distinct long-term climatological gradientsof temperature and light. The 1960s were generally colder andbrighter than the 1970s. Prior to 1969, the annual phytoplanktonmaximum occurred most commonly during winter; in the 1970s,the annual maximum generally shifted to a summer event. Three5-year phytoplankton cycles occurred between 1959 and 1974.During each pentade, the phytoplankton community returned toa similar taxonomic organization and abundance cycle after divergingin the intervening years. Pentade cycles did not occur after1974; the phytoplankton community thereafter diverged significantlyfrom each preceding year. Five species [Skeletonema costatum,Detonula confervacea, Asterionellopsis glacialis, Hererosigmaakashiwo (= Olisthodiscus lureus) and Thalassiosira nordenskioeldii]dominated the phytoplankton over the 22-year period. SDA revealeda high degree of similarity and constancy in the annual occurrencepatterns of these taxa. The decadal shifts revealed by SDA weremore directly related to the considerable interannual variabilitythat characterized the abundance and seasonality of the lessabundant species.     

Export of Nitrogen,Phosphorus, and Suspended Solids from a Southern New England Watershed to Little Narragansett Bay

The Pawcatuck River watershed (764 km²) is a mainly forested drainage basin with a low population density (80 people km⁻²) that discharges to a shallow estuary, Little Narragansett Bay (RI and CT, USA). In order to quantify the nitrogen (N) and phosphorus (P) flux to the estuary, we measured all forms of nitrogen and phosphorus, as well as suspended solids at the mouth of the river above tidal influence, on more than 80 occasions over an annual cycle. The annual export of total nitrogen, total phosphorus, and total suspended solids amounted to 16.0×10⁶ mol y⁻¹, 0.97×10⁶ mol y⁻¹, and 1.4×10⁶ kg y⁻¹, respectively. Nitrogen export was equally divided between dissolved inorganic (83% NO₃⁻) and organic forms, with particulate nitrogen comprising 17% of the total flux. Phosphorus export was dominated by particulate forms (67%), with dissolved inorganic phosphate contributing 30% and dissolved organic phosphorus contributing 8% of the annual flux. Preliminary nutrient budgets for the Pawcatuck watershed suggest that only about 10% of the nitrogen and phosphorus inputs are exported from the system. Strong regressions between water discharge and TN enabled us to extrapolate the data collected during the relatively dry study period to a long term average discharge year. Under normal river discharge conditions, the N flux would be approximately 26.0×10⁶ mol y⁻¹ or about 20% of the nitrogen inputs to the watershed. This value is very close to the N flux predicted by a regression developed by others from a wide range of larger watersheds. The relatively large size of the Pawcatuck watershed relative to the estuary (9.6 km²), makes Little Narragansett Bay one of the most intensively nitrogen loaded estuaries on the Atlantic coast in spite of the dominant forest cover of the watershed.     

Seasonal fluctuation of zooplankton populations in lower Delaware Bay

Quantitative zooplankton samples were obtained monthly or bi-monthly 15 times from June 1974 to May 1975 at three stations in lower Delaware Bay. Two 12-hour cruises were also conducted at one of the stations.Arthropods dominated the samples in terms of number of species and number of individuals. The number of zooplankton from surface samples ranged from 58/m³ in August to 21,092/ m³ in June, while bottom samples varied from 259/m³ in August to 30,395/ m³ in October. In general, larger concentrations of individuals were found in bottom samples.Only on three occasions did meroplankton exceed the holoplankton, and these occurred at the shallow water stations. Meroplankton comprised a larger percentage of the bottom samples than surface samples. Zoeae of Neopanope sayi and Uca sp. contributed mainly to the large proportion of meroplankton in July 1974, veligers of Mytilus edulis in January 1975, and nauplii of Balanus sp. in May 1975.Copepods were the largest component of the population throughout most of the year. At all stations and depths, Arctica tonsa dominated most of the summer samples. In the spring of 1975, A. tonsa was replaced by Centropages hamatus, Temora longicornis, and Pseudocalanus minutus.During the 12-hour cruises there were higher numbers of individuals in the bottom waters in the day with migration to surface waters in the afternoon and evening. Based on cluster analysis, five time-related assemblages were discerned: June, July–August, September–November, December, January–May. Comparison of Delaware Bay zooplankton with other estuarine systems indicated that the densities obtained locally were most similar to those reported in the York River, Virginia.     

Annual phytoplankton cycle in a meromictic anoxic basin of a Rhode Island (USA) estuarine river

Hydrobiologia - Estuarine Pettaquamscutt River is a unique habitat 10&nbsp;km in length with physical and biogeochemical characteristics analogous to a miniature fjord. Its meromictic upper...     

Evaluation of sediment profile imagery as a tool for assessing water quality in Greenwich Bay,Rhode Island,USA

The Benthic Habitat Quality (BHQ) index was used to assess habitat visible in sediment profile images (SPI) following hypoxia disturbance in a shallow (<10 m) estuarine embayment in Rhode Island, USA. We tested for associations between the BHQ, SPI features and water quality over several assessment windows (1, 3, 7, 14, and 28 days prior to imaging) and at multiple dissolved oxygen (DO) thresholds (2.0 mg l^?1, 2.9 mg l^?1, and 4.8 mg l^?1). Using categorical data analysis, we established empirical relationships between hypoxia prevalence and presence/absence of biogenic features visible in SPI. Fecal pellets, tubes, feeding pits, voids, mounds, and BHQ score were good affirmative features, meaning that their presence (or score greater than 5) indicated a high probability of good water quality. However, low sensitivity to hypoxia precluded their usefulness as indicators, and was attributed to rarity in images and to factors acting on time intervals longer than those examined, e.g. long-term organic enrichment or hypoxia. Burrow structures and the apparent redox potential discontinuity (aRPD), or oxidized layer of surface sediment, were good discriminatory features, with high sensitivity and specificity for both hypoxia and normoxia. Both were strong surrogates for water quality over multiple assessment windows and DO thresholds, and had the highest overall predictive values. We conclude that SPI images can be used to widen the spatial extent of water quality monitoring efforts by utilizing the relationships between aRPD, burrows and hypoxia prevalence.     

Underestimation of primary production as indicated by measurements with size-fractionated phytoplankton in Lake Maarsseveen I (The Netherlands)

Primary productivity of four size classes of phytoplankton (<150m, <50m, <20m and <5m) was measured from March through October 1986 in Lake Maarsseveen I with an incubator technique. The mean column production was approximately 400 mg C.m^–2.day^–1, with a range of values between 150 and 750 mg C.m^–2.day^–1. The mean contribution of the size fractions <50m, <20m and <5m to the size fraction<150m was 80%, 60% and 35%, respectively. During their appearance the grazing impact of small herbivorous zooplankton,e.g. rotifers, can give an underestimation of the size fraction <150m. An indication of this phenomenon is given.     

Studies on the phytoplankton ecology of the Trondheimsfjord. I. The chemical composition of phytoplankton populations

Samples of phytoplankton populations from the Trondheimsfjord, collected in 1970 and the first five months of 1971, have been analysed for carbohydrate, protein, lipid, and phosphorus. Lipid was in all cases less than 10% of the organic dry matter. The $\text{N}\text{P}$ ratio was remarkably constant, but the ratio protein/carbohydrate varied between wide limits. For samples consisting mainly of dinoflagellates, the protein/carbohydrate ratio was always low, due to a large amount of insoluble polysaccharides, probably corresponding to material in the cell walls.For diatoms, the carbohydrates may conveniently be divided into three fractions: 1) an acidsoluble glucan of the β-1, 3-linked type; 2) an alkali-soluble fraction giving a complex mixture of monosaccharides on hydrolysis and, 3) an insoluble glucan. The amounts of acid-soluble glucan varied from 7.7 to 36.5% of organic dry matter and these changes are the main cause of the variation of the protein/carbohydrate ratio of diatom samples. For diatom samples this ratio is a valuable indicator of the physiological state of the population. The variations observed in this study are discussed.     

Nitrogen uptake by size-fractionated plankton in permanently well-mixed temperate coastal waters

Nitrogen uptake by net- (15–200 µm), nano- (1–15µm) and picoplankton (<1 µm) was measured overseasonal cycles at two stations with different patterns of biologicaland chemical cycles in the Morlaix Bay (western English Channel).Though assimilable dissolved N nutrient pool at both stationswas nitrate-dominated, characteristics of biomass and N uptakeby netplankton differed from conventional patterns in two respects.In the first, biomass (26–30%) and N uptake (36–43%)were less important than those of nanoplankton. In the second,the netplankton did not show any marked preference for nitrateover ammonium (nitrate to ammonium uptake ratios of 0.98 and1.08). In contrast, nanoplankton had a preference for ammoniumover nitrate (ammonium to nitrate uptake ratios of 2 and 1.2).N uptake by picoplankton was only 8% of total N uptake at bothstations and was supported mainly by regenerated N (66% ammoniumand 17% urea), with nitrate uptake detectable in only one instanceand nitrite uptake in none. Substrate-dependent uptake of ammoniumin all fractions and a higher ammonium uptake in the nanoplanktonfraction in summer at both stations when ambient ammonium concentrationswere high indicated that while nitrate may satisfy a part ofN requirements, availability of ammonium and its flux throughnanoplankton determine the magnitude of total N uptake in thesewaters. Most of the N uptake in picoplankton appears to be autotrophic,suggesting that a substantial part of heterotrophic uptake,if any, could be localized in the fractions >1 µm,and mediated by free-living and particle-bound bacteria.     

Growth, production and losses of phytoplankton in the lowland River Spree. I. Population dynamics

The seasonal and longitudinal dynamics of dominant phytoplanktonpopulations were observed in the lower Spree (Germany), a lowlandriver draining several lakes. Centric diatoms and cyanobacteriawere the chief components of the total phytoplankton biomassof 16 mg^–1 at the end of the river course (average March-October,1988–89). The cyanobacteria required a retention timeof several weeks in a drained lake to establish stable populations,whereas the other algal populations originated from upstreamimpoundments and slow-flowing river stretches. In spring, thedominant populations increased (the diatoms Nitzschia acicularis,Asterionella formosa, Stephanodiscus hantzschii. Cyclostephanosdubius, Cyclotella radiosa, C.meneghiniana) or kept their abundancesnearly constant (cyanobacterium Limnothrix redekei. diatom Synedraacus), but the biomass of the ‘summer species’ (diatomActinocyclus normanii, cyanobacteria Planktothrix agardhii andAphanizomenonsp.) was halved along the river course. These seasonal differencesin algal growth are probably caused by the changed mixing patternin the regions of origin and thus different pre-adaptation toturbulent mixing. Pronounced dear-water phases occurred regularlyat the end of May. They were intensified along the river coursedespite the lack of mesozooplankton. The dynamics of phytoplanktonwere reflected in the proportions of dissolved and particulatenutrients. The nutrient supply was always high enough to makelimitations of algal growth improbable.