A 91-year record of seasonal and interannual variability of diatoms from laminated sediments in Saanich Inlet, British Columbia

Diatom seasonal succession and interannual variability werestudied using laminated sediments from Saanich Inlet, BritishColumbia, for the years 1900–1991. Frozen sediment coresallowed fine-scale sampling of laminae for each year. Thus,three ‘seasons’ for each year were identified basedon species composition. Thalassiosira species were indicatorsof spring deposition. Skeletonema costatum was abundant in samplesfollowing Thalassiosira, probably deposited in late spring andsummer. Rhizosolenia sp. was most abundant in fall/winter samples.Diatom stratigraphies were related to sea surface temperature,salinity, sea level and the Pacific North American Index (PNA)using canonical correspondence analysis (CCA). CCA showed thatspecies of a particular season generally had optima for temperatureand salinity characteristic of that time. Interannual changesin diatom species composition and abundance were most prevalentin the decades 1920–1940, with the exception of S.costatumwhich showed cyclic changes in abundance. Skeletonema was moreabundant during periods of cool temperatures, while littoraldiatoms were more abundant during times of heavy winter rains.Sea level was an important variable in CCA and while its relationshipto diatoms is not clear, it may be related to variations innutrient supply to diatoms in surface waters.     

Fluxes of diatoms in the Dona Paula Bay, west coast of India

Sediment traps were deployed at a station in the Dona PaulaBay to collect sedimenting particles at weekly intervals fromNovember to May during 1995–1997. Sedimented particleswere analysed for total diatom flux, chlorophyll a (Chl a) andparticulate organic carbon (POC). The highest diatom flux wasrecorded in April–May for both the years. Fluxes of diatomsvaried from0.6 x 10⁴ cells m^–2 day^–1 (November 1995)to 121.47 x 10⁴ cells m^–2 day^–1 (December 1996).In all, 19 diatom genera were identified in the sedimented material.Navicula, Nitzschia, Pleurosigma, Licmophora, Coscinodiscus,Rhizosolenia and Surirella were the most abundant genera inthe sedimented material throughout the sampling period. Meanflux of POC and diatom carbon was 251 and 0.39 mg C m^–2day^–1, respectively. The diatom carbon accounted for 0.15%of the POC flux. Mass flux of diatoms showed significant negativecorrelation with the concentration of nitrate and phosphate.This suggests that the nutrient concentration played an importantrole in influencing the sedimentation of diatoms at this coastalstation.     

Infection of Coscinodiscus spp. by the parasitoid nanoflagellate Pirsonla diadema: I. Behavioural studies on the infection process

The investigation of successive steps involved in the infectionprocess of the marine diatoms Coscinodiscus granii and Coscuwdiscuswailesii by the host-specific parasitoid nanoflagellate (PNF)Pirsonia diadenw showed that flagellates reacted chemokJinokineticallywith changes of swimming pattern to the presence of a host diatom.Chemosensory stimulation appeared to induce readiness for infection,whereas attachment and penetration of the diatom cell wall wasinduced by a mechanosensory response to morphological featureson the diatom frustules. The mean swimming speed of P.diademaflagellates decreased during their infective lifetime of 3 daysfrom an average of 78 µm s^–1 to 51 µm s^–1while the frequency of small loops in the swimming pattern increasedfrom 0.8 to 6.3 loops min^–1. At high Cgranii densities,an epidemic was delayed. It is suggested that this could becaused by overlapping gradients of extracellular material releasedby the diatoms which impaired the sensing of spatial gradientsby PNF and, therefore, the location of hosts.     

Infection of Coscinodiscus granii by the parasitoid nanoflagellate Pirsonia diadema: III. Effects of turbulence on the incidence of infection

The influence of turbulence on the incidence of infection ofthe diatom Coscinodiscus granii by the parasitoid nanoflagellatePirsonia diadema was investigated experimentally with two initialhost densities. Independently of the initial diatom densitiesof 7 and 44 cells ml^–1, under calm conditions both diatomsand parasitoids became extinct within 6–9 days. Turbulence,however, led to the survival of diatoms at a reduced densityof 0.1–2 cells ml^–1 for >30 days. A population-dynamicmodel is formulated that takes into account the non-homogeneousdistribution of infecting flagellates among host diatoms. Applicationof the results to parasitoid–diatom interactions in naturalwaters suggests that, under turbulent conditions, endemic infectionsmay effectively prevent the mass development of host diatoms.     

Studies in the Diatom Genus Hantzschia 3. Infraspecific Variation in H. virgata

Infraspecific variation has been investigated in the marineepipelic diatom Hantzschia virgata using light and electronmicroscopy. Although the variation pattern in this species iscomplex and to some extent discontinuous, allowing the recognitionon phenetic grounds of four or five discrete infraspecific taxaincluding vars. virgata (= H. insolita), gracilis, leptocephala(= capitellata) and wittii (= intermedia), it is concluded thatto split the species would be imprudent. Variation has beenfound not only in characters such as valve outline and size,or stria density, but also in ‘qualitative’ characterssuch as raphe, fibula and girdle structure, characters whichhave often been assumed to be constant within species, generaor taxa of higher rank: previous generalizations about the usefulnessof certain characters in distinguishing taxa of a given rankare seen to have been incautious. Hantzschia virgata (Roper) Grun., diatom taxonomy, infraspecific variation     

Experimental studies of food and suspended sediment effects on growth and reproduction of six planktonic cladocerans

Life-long somatic growth and egg production responses of Daphniabarbata, Daphnia gibba, Daphnia laevis, Daphnia pulex, Moinamicrura and Diaphanosoma excisum were examined in relation todifferent food and suspended sediment levels, in factorial experimentsat 20°C. Food treatments involved additions of 0, 100, 500and 2500 µg C l^–1 of Selenastrum to 20 µmfiltered water, generally from source lakes. These food levelswere tested at two suspended sediment levels in all species,viz. (i) natural source-lake nephelometric turbidities of 7–65nephelometric turbidity units (NTU) ({small tilde}10–70mg l^–1 TSS) and (ii) {small tilde}3- to 4-fold enrichmentof natural levels (8-fold in D.excisum). A third sediment-freetreatment was tested in all species of Daphnia other than D.gibba.Growth in body length and reproductive output were assesseddaily for individual animals. Main and interactive effects offood and sediment on growth were assessed at selected ages fromday 3 through subsequent life using analysis of variance (ANOVA).Strong and highly significant positive food effects were apparentat all ages in all species examined, with the curious exceptionof D.excisum. The onset of maturity and fecundity were alsopositively related to food level. Sediment effects on growth,age of maturity and fecundity were variable, but tended to bemuch weaker than food effects. Both stimulatory (positive, especiallyat low food levels) and inhibitory (negative) effects of sedimenton growth were apparent, both between and within species. Performancein sediment-free water was very poor, except in D.pulex. Food-sedimentinteraction effects were variable. Generally, turbidwater specieswere influenced less adversely by sediment than clear-waterspecies, indicating the existence of environmentally appropriateadaptive responses. Overall, the findings indicate that lightlimitation of algal resources by suspended sediments, ratherthan the condition of high turbidity per se, is responsiblefor the apparently negative impact of suspended sediments frequentlyobserved on natural populations of cladoceran zooplankters.     

Diatom preservation: experiments and observations on dissolution and breakage in modern and fossil material

Selected aspects of diatom preservation in both laboratory and field environments are examined with a view to improving techniques and to help understand why only some lake sediments have good diatom preservation.Laboratory measurements of biogenic silica following diatom dissolution by alkali digestion are questioned because results are shown to be dependant on initial sample size. Diatom breakage experiments identified drying carbonate rich sediment as a major cause of fragmentation of the large robust diatom Campylodiscus clypeus Ehrenb. Diatom dissolution experiments in carbonate media indicated that carbonate rich lakes should preserve diatoms better in order of the particular alkali metal type (Ca > Mg > Na). A preliminary assessment of the role of depth in diatom preservation is made for Lake Baikal where partly dissolved Cyclotella are more common in deep water surface sediments. The effect of time on diatom dissolution is examined in a saline lake sediment core and by comparing dissolution rates of recent and geologically old diatom samples in the laboratory. A simple link between diatom dissolution and sample age was not established. Factors thought to be important in controlling diatom preservation in lake sediments are discussed.     

A Study of Two Morphological Variants of the Diatom Fragilaria crotonensis Kitton using Electron Microscopy

Two forms of Fragilaria crotonensis Kitton, one with flaredends to the cells, the other with ends that taper to a narrowrod-like tip, were previously distinguished by light microscopyand by their contrasting relationships with the parasitic chytridsRhizophydium fragilariae Canter and one referred to as species3. Examples of the above diatom types, taken from a number ofgeographically widely separated lakes, have been examined byelectron microscopy. Additional morphological differences aredescribed. The ocellus in flared-type cells consists of 4–5rows of pores while in rod-type cells fewer (2–3) rowsof pores are present. Valve thickenings on the mantle at theapex of rod-type cells are rarely visible whereas in flared-typecells they are usually very prominent. A rimoportula was presentin both cell types. In flared cells its external aperture liesimmediately behind the apex usually in line with the striaeand the internal aperture is often orientated with its longaxis oblique to that of the diatom cell. The rimoportula inrod-type cells opens at the centre of the valve face and theinternal aperture is orientated at 90° to the apical axisof the valve. Comparison made with type material suggests thatthe rod-form should be recognized as F. crotonensis var. crotonensisand the flared-form as var. prolongata Grunow ex van Heurck. Fragilaria crotonensis, diatom, systematics, chytrid infection, rimoportula     

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.     

Horizontal distribution of Thetys vagina Tilesius (Tunicata, Thaliacea) in the Japan Sea during spring 2004

The occurrence of the salp Thetys vagina was observed in theJapan Sea during spring 2004. Catches up to 187 kg wet weight(WW) per 2.18 x 10⁵ m³ (equal to 0.9 g WW m^–3) were collectedwith 10-m diameter surface-water otter trawl nets. The horizontaldistribution indicated that the high biomass was related tothe area with high chlorophyll a (Chl a) concentration, whichwas located around the subarctic front with the warm TsushimaCurrent. Five prey taxa were identified from the gut contentsof individuals from the high Chl a area. The diatom Coscinodiscusspp. (13–55 µm in diameter) dominated numerically.Another significant prey was the large diatom Coscinodiscuswailesii (219–313 µm) that is an indicator of thespring bloom in this area. The mass occurrence of T. vaginathus appears related to phytoplankton availability, though themechanisms remain uncertain.     

Diatom assemblages in sediments of Lake Juusa, Southern Estonia with an assessment of their habitat

Diatom analysis of surface sediments and two sediment cores from different sedimentation areas of a small closed lake was undertaken with the aim of acquiring knowledge on the dependence of the distribution of diatom assemblages on lake bathymetry. Lake Juusa was selected for the study because we have for this lake a large data set about the lithological composition of sediments and macrofossil and cladoceran records for the Holocene. A high carbonate content (20–60%) in the sediment sequence indicates high carbonacity and relatively stable pH values during the Holocene. On the basis of comprehensive analysis, abrupt water-level fluctuations and changes in the trophic status were established. Results of this study showed that the fluctuations of the water-level were the leading factor determining the habitats of diatom assemblages in the lake. In the surface sediment samples planktonic species such as Cyclotella spp., Stephanodiscus spp. and Aulacoseira spp. had a depth optimum at 3–4 m and the most abundant periphytic taxa were distributed mostly at depths shallower than 3.5 m. The same regularity was established in sediment cores where a good correlation between planktonic species and lake water depth was found in sediments accumulated at water depths >4 m. Lake Juusa appears to be a proper site for detailed environmental reconstructions over the Holocene, and the results will give us a good opportunity to analyse the history of water-level fluctuations in other small Estonian lakes. Electronic Supplementary Material The online version of this article (doi:) contains supplementary material, which is available to authorized users. Handling editor: K. Martens     

Occurrence of viable photoautotrophic picoplankton in the aphotic zone of Lake Biwa, Japan

The distribution and abundance of photoautotrophic picoplankton(PPP. Synechococcus group) in the aphotic bottom sediments ofLake Biwa were investigated by direct counting and viable counting(most probable number, MPN) methods. In the surface layer ofbottom sediments (0–1 cm). where large PPP blooms occurredin the past 5 years, >10⁵ cells cm^–3 of PPP were foundto be viable throughout the year. Furthermore, the density ofPPP deposited on the sediment surface (0–0.1 cm) was oneorder of magnitude higher (MPN = 1.3 x 10⁶ cells cm^–3.direct count = 9.9 x 10⁶ cells cm^–3) than that of bulkedsurface sediments (0–1 cm). Even in the deeper layer (13–14cm) of bottom mud, viable PPP were still found (10¹ cells cm^–1.In winter, viable PPP in the aphotic bottom sediments were 10⁴–10⁵times greater per Unit volume than those in the euphotic lakewater. Since the aphotic bottom sediments have high levels ofPPP, as well as high growth potential (high ratio of viablecount/total direct count), they are likely to seed PPP bloomsin the North Basin of Lake Biwa.     

Increase in silicon concentrations and release from suspended solids and bottom sediments in Lake Kasumigaura, Japan

Increasing trends of dissolved Si measured by a colorimetric method and ICP (DSi_col and DSi_ICP, respectively) and total Si concentrations were detected at the center of Lake Kasumigaura during 1980–2006 (mean DSi_col concentration in the 1980s and 2000s was 1.3 and 4.0 mg l⁻¹, respectively). The observation of such trends is rare; therefore, the elucidation of the causes could be useful to understanding silicon dynamics in inland waters. Based on statistical analysis, we found that the increases in DSi_col and lithogenic Si accounted for most of the total Si increase (44 and 45%, respectively) and that biogenic Si, consisting of diatom frustules, also increased with them. Increases in DSi_ICP concentration were not detected near the mouth of the inflowing rivers, suggesting that the increase was caused by in-lake processes. Because the increase in suspended solids (SS) caused by sediment resuspension had been observed in the lake for the same period, we assumed that the Si release from SS containing diatom frustules contributed to the increase. The results of the laboratory experiments in which surface sediments were stirred in lake waters showed that the change in DSi_col concentration depended mainly on SS concentration, water temperature and the elapsed time of diatom frustules dissolution. An estimation of the released amount of Si from SS using the sediment resuspension model was (1.0–2.7) × 10⁹ g year⁻¹ in the 2000s, which was about 30–90% of the increase in the DSi_col outflow of 3.0 × 10⁹ g year⁻¹ from the 1980s to the 2000s. We also determined the Si release rates from bottom sediments through laboratory experiments. The Si amount released from bottom sediments in the lake in the 2000s was estimated to be 4.3 × 10⁹ g year⁻¹, which was about 2–4 times higher than the estimated Si amount released from SS. These findings suggest that the sediment resuspension might be the cause of the latest DSi increase.     

Sinking losses of phytoplankton in closed limnetic systems

Specific algal recoveries from sediment traps of two differentdesigns and from mud surface deposits of large experimentalenclosures (Lund Tubes) were monitored during 1978 and are analyzedin relation to the vertical and temporal distribution of tendominant phytoplankton populations. Sedimentation accounts fordiffering proportions of the total loss of biomass for differentalgae: between 28 and 100% of diatoms; 15–95% of Eudorina;<4% of populations of small algae (spp. ofAnkyra, Chromulina,Cryptomonas). Rates of diatom loss are also derived from thecomparison of net rates of change (k_n) and the silica uptake-derivedgrowth rate (k¹); intrinsic sinking behaviour may be specificallyregulated in relation to growth conditions. Implications inthe calculation of sedimentary losses and their impact uponthe seasonal periodicity of phytoplankton are briefly discussed.     

Seasonal and microhabitat influences on diatom assemblages and their representation in sediment traps and surface sediments from adjacent High Arctic lakes: Cape Bounty,Melville Island,Nunavut

The spatial (i.e. microhabitat) and temporal (i.e. seasonal) characteristics of diatom assemblages in adjacent High Arctic lakes were studied intensively June–August 2004. These baseline data are used to improve understanding of modern diatom community dynamics, as well to inform paleoenvironmental reconstructions. Diatoms were collected approximately weekly through the melt season from each principal benthic substrate (moss/macrophyte, rock scrapes, littoral sediment), plankton, and sediment traps, and were compared to the uppermost 0.5 cm of a surface core obtained from the deepest part of the lake where sediment cores are routinely collected. Water samples were collected concurrently with diatom samples to investigate species–environment relationships. The lakes share approximately half of their common taxa, the most abundant overall in both lakes being small Cyclotella species. Results of detrended correspondence analysis (DCA) indicate that the largest gradient in species turnover existed between benthic and planktonic communities in both lakes, and that sediment trap and the surface core top samples most closely resemble the planktonic assemblage, with an additional contribution from the lotic environment. Our results indicate clear micro-spatial controls on species assemblages and a degree of disconnection between the benthos and deep lake sediments that manifests as an under-representation of benthic taxa in deep lake surface sediments. These findings are particularly relevant in the context of interpreting the paleoenvironmental record and assessing ecosystem sensitivity to continued climate change.     

A change in the distribution pattern of diatoms above and below the halocline in bottom sediments of the central Arkona Basin,southern Baltic

Surface sediment samples from the central Arkona Basin, southern Baltic, were investigated by diatom analysis. The samples, taken by a Niemistö sampler, included the sediment-water interface and about 2–3 centimetres of the very water-saturated sediment. The samples represent the sediments deposited in the transition layer of the halocline in the Arkona Basin and immediately above and beneath it. The distribution pattern of diatoms and diatom pieces changed, reflecting the position of the halocline.Diatom analysis is an important tool in identifying Holocene Baltic sediments, and the knowledge of recent hydrological and chemical factors and their impact on the distribution pattern of diatoms is one way to evaluate the usefulness of diatom records in postglacial Baltic marine sediments. The surface-sediment diatom assemblages can be compared with the data from postglacial Baltic offshore cores. They give prelimary indications where diatom records from Littorina or Yoldia stages could be expected.     

Changes in the growth rates of saline-lake diatoms in response to variation in salinity, brine type and nitrogen form

The growth rates of four saline-lake diatom taxa were measuredunder varying conditions of salinity (5, 8 and 11), brine type(sulfate- versus bicarbonate-dominated) and nitrogen form (NH₄⁺versus NO₃^–), using a full factorial design. With NO₃^–as the nitrogen source, Cyclotella quillensis, Cymbella pusillaand Anomoeoneis costata exhibited lower growth rates in thesulfate versus bicarbonate media. The strain of Chaetoceroselmorei used in these experiments, isolated from a sulfate-dominatedlake, was unable to grow on NO₃^– alone. In the NH₄⁺ treatments,neither salinity nor brine type affected the growth rates ofC.quillensis or C.elmorei. When supplied with NH₄⁺, C.pusillaand A.costata had higher growth rates in the bicarbonate versussulfate media, although for C.pusilla the difference on NH₄⁺was not as great as on NO₃^–. The impact of brine typeon NO₃^– use is consistent with the theory that sulfateinhibits molybdate uptake, as molybdenum is required for NO₃^–use but not NH₄⁺. Cymbella pusilla was the only taxon affectedby changes in salinity. The four taxa used in these experimentsare frequently found in saline lakes and saline-lake sediments,hence they are used in paleoclimate reconstructions; the resultspresented here provide additional information that may enhancethese diatom-based reconstructions.     

Diatom taphocoenoses in the coastal upwelling area off South West Africa

The diatom floral composition of 124 sediment samples from the South East Atlantic records the influence of coastal upwelling on sediment composition off South West Africa. Inner shelf samples between 19° and 24°S are rich in diatom valves and the patterns of diatom species distribution in these samples are related to the coastal upwelling process. Comparison with recent phytoplankton data shows that the sediment assemblages preserve many of the important species of the diatom biocoenoses, including Chaetoceros (resting spores), Delphineis karstenii, Thalassiosira eccentrica, and Thalassionema nitzschioides. Delphineis karstenii, a pioneer species in enriched coastal water, occurs nearshore and Chaetoreros resting spores are widespread, with highest relative abundance values in some more offshore samples. The abundance of the Thalassiosira eccentrica group and of Thalassionema nitzschioides in sediment samples in and near Walvis Bay reflects the recurrence of intense upwelling off this part of the coast. An abundance of large centric species has been reported in hydrological conditions characteristic of newly upwelled waters and, correspondingly, Actinocyclus octonarius and some large Coscinodiscus species occur in the sediments in nearshore patches or belts.     

Effects of concentration on the feeding of a marine copepod in algal monocultures and mixtures

High-speed microcinematography was used to examine the feedingbehavior of the marine copepod Eucalanus elongatus in a rangeof concentrations of algal monocultures and mixtures. Two celltypes were offered, the 13-µm diatom Thalassiosira weissflogii,which is primarily accumulated passively by low amplitude flappingof the second maxillae, and the 20 450-µm diatom Rhizosoleniaalata, which is actively captured by detection of and orientedresponse to individual cells. E. elongatus rapidly switchedback and forth between these two capture modes in mixtures ofboth diatoms, and flapped the second maxillae at low amplitudesregardless of the absolute or relative abundance of small andlarge cells. However, copepods in both monocultures and mixturesaltered the duration and/or rate of flapping of the feedingappendages with changes in algal concentration, with maximumactivity levels occurring at intermediate concentrations. Themarked reduction in feeding motions observed at the lowest algalconcentrations supports results from traditional grazing studiesand optimal foraging models, and may conserve energy duringprolonged perods of low food availability in continental slopewaters.     

In situ egg production and hatching success of the marine copepod Pseudocalanus newmani in Funka Bay and adjacent waters off southwestern Hokkaido, Japan: associated to diatom bloom

. Clutch size and egg viability were examined for wild femalesof Pseudocalanus newmani collected from four different stationsinside and outside Funka Bay, southwestern Hokkaido, Japan,during pre-phytoplankton-bloom conditions in 1997, and froma single station outside the bay during the period from pre-to post-blooming seasons in 1998, to examine spatio-temporalvariability of the reproductive parameters and to detect deleteriouseffects of diatom blooms on in situ copepod reproduction. Clutchsize and hatching success in P.newmani were relatively similarat the four stations in 1997. The eggs almost always hatchedcompletely, but deformed nauplii occurred in 20–40% ofthe cases. The proportion of deformed nauplii gradually decreasedwhen the females were fed on non-diatom Pavlova sp. in the laboratory,suggesting that the occurrence of deformed nauplii was relatedto the quality of in situ food particles, including diatoms.Clutch size varied from 12 to 30 eggs female^–1 duringthe study period in 1998; more than 90% of the variation couldbe explained by the body size of the females which dependedon ambient water temperature. Hatching success also varied,from 15 to 80%, but was not related to either clutch size, femalebody size, water temperature or diatom biomass. These resultssuggest that not only egg production but also hatchability shouldbe measured routinely when estimating recruitment of the copepodinto the planktonic population, and show that neither clutchsize nor egg viability are directly affected by diatom biomass.Although we could not obtain clear evidence of in situ deleteriouseffects of diatoms on abnormal embryos and nauplii, this mightbe related to methodological problems and properties or characteristicsof prey–predator interactions. Also, deformities of copepodnauplii have not yet been observed for reasons other than feedingon diatoms.