ASSOCIATION OF A FREE-LIVING EUGLENA WITH A COLONIAL ROTIFER

Large epiphytic and epilithic diatom species hosting other diatoms were observed in several fresh- and brackish water sites in Southern California. The most commonly encountered hosts were species forming long filaments attached to rocks or macroalgae, Hydrosera whampoensis (Schwarz) Hendey, Melosira varians Agardh, Pleurosira laevis (Ehrenberg) Compere and Terpsinoe musica Ehrenberg. These large diatoms often had smaller diatoms attached, usually to the girdle bands and occasionally to the mucilage pads connecting the cells. For example, cells of T. musica were observed supporting growth of a diverse diatom assemblage composed of species of the genera Achnanthes , Achnanthidium , Amphora , Cocconeis and Tabularia; Synedra sp. was attached to M. varians and B. paxillifer; and Cocconeis placentula was seen on H. whampoensis. Thus, large epiphytic and epilithic diatoms seem to provide suitable sites for attachment of small epiphytic diatom species, and it appears that this phenomenon is more common than previously thought.     

DIATOMS AS HOSTS FOR OTHER DIATOMS: OBSERVATIONS FROM SOUTHERN CALIFORNIA

Large epiphytic and epilithic diatom species hosting other diatoms were observed in several fresh‐ and brackish water sites in Southern California. The most commonly encountered hosts were species forming long filaments attached to rocks or macroalgae, Hydrosera whampoensis (Schwarz) Hendey, Melosira varians Agardh, Pleurosira laevis (Ehrenberg) Compere and Terpsinoe musica Ehrenberg. These large diatoms often had smaller diatoms attached, usually to the girdle bands and occasionally to the mucilage pads connecting the cells. For example, cells of T. musica were observed supporting growth of a diverse diatom assemblage composed of species of the genera Achnanthes, Achnanthidium, Amphora, Cocconeis and Tabularia; Synedra sp. was attached to M. varians and B. paxillifer; and Cocconeis placentula was seen on H. whampoensis. Thus, large epiphytic and epilithic diatoms seem to provide suitable sites for attachment of small epiphytic diatom species, and it appears that this phenomenon is more common than previously thought.     

COASTAL DIATOM‐ENVIRONMENT RELATIONSHIPS IN THE BRACKISH BALTIC SEA1

High‐quality calibration data sets are required when diatom assemblages are used for monitoring ecological change or reconstructing palaeo‐environments. The quality of such data sets can be validated, in addition to other criteria, by the percentage of significant unimodal species responses as a measure of the length of an environmental gradient. This study presents diatom‐environment relationships analyzed from a robust data set of diatom communities living on submerged stones along a 2,000 km long coastline in the Baltic Sea area, including 524 samples taken at 135 sites and covering a salinity gradient from 0.4 to 11.4. Altogether, 487 diatom taxa belonging to 102 genera were recorded. Detrended canonical correspondence analysis showed that salinity was the overriding environmental factor regulating diatom community composition, while exposure to wave action and nutrient concentrations were of secondary importance. Modeling the abundances of the 58 most common diatom taxa yielded significant relationships with salinity for 57 taxa. Twenty‐three taxa showing monotonic responses were species with optimum distributions in freshwater or marine waters. Thirty‐four taxa showing unimodal responses were brackish‐water species with maximum distributions at different salinities. Separate analyses for small (cell biovolume <1,000 μm³) and large (≥1,000 μm³) taxa yielded similar results. In previous studies along shorter salinity gradients, large and small epilithic diatom taxa responded differently. From our large data, we conclude that counts of large diatom taxa alone seem sufficient for indicating salinity changes in coastal environments with high precision.     

Uncertainty of using habitat fidelity in biomonitoring based on benthic diatoms - the Raška River case study

Epilithic diatoms are frequently recommended for river biomonitoring, while much less emphasis is placed on epiphytic communities. This study considers use of epiphytic and epilithic diatom communities from the Ra?ka River in biomonitoring. A total of 212 diatom taxa were recorded in both communities. Dominant diatoms were Achnanthidium minutissimum (Kützing) Czarnecki, A. affine (Grunow) Czarnecki, A. subatomus (Hustedt) Lange-Bertalot, Amphora pediculus (Kützing) Grunow, Diatoma vulgaris Bory, Gomphonema tergestinum (Grunow) Fricke, Cocconeis placentula var. lineata (Ehrenberg) Van Heurck, Melosira varians Agardh and Navicula tripunctata (Müller) Bory. Redundancy analysis (RDA) was used to detect the relationships between diatoms, some environmental factors and sampling sites. Diatoms of the epiphytic community showed a clearly grouping in relation to the sampling sites as compared with diatoms of the epilithic community. Species common to both communities showed a similar correlation with some environmental variables (BOD, pH, NH₄⁺, TP, NO₃^? and conductivity), with three exceptions (Diatoma vulgaris, Cymbella compacta, and Encyonema silesiacum). Pearson’s correlation coefficient showed correlation between selected environmental variables and diatom indices calculated using OMNIDIA 6.2 software (TDI, IPS, CEE, GENRE, TID, SID, SHE and IDSE/5). Diatoms of the epiphytic community are more clearly clustered in relation to the sampling sites as compared with diatoms of the epilithic community. According to our results there is still “place” in biomonitoring for epiphytic community. The present study highlights the necessity of sampling both epiphytic and epilithic communities because substrate specificity could play important role in biomonitoring.     

Photosynthetic activity of benthic diatoms in response to different temperatures

The photosynthetic activities of benthic diatoms in response to temperature changes were assessed by measuring chlorophyll fluorescence kinetics. Small benthic diatom species with large surface area to volume (SA/V) ratios responded to increasing temperature differently from large diatoms, since larger ratios caused lower photosynthetic activity under high-temperature conditions. The small SA/V ratios of large cells may be advantageous in benthic environments under adverse conditions such as high temperature and/or strong light. A size-dependent differential response of benthic diatoms to changes in environmental factors such as temperature may result in an altered distribution of the different diatom communities.     

Stream epilithic, epipelic and epiphytic diatoms: habitat fidelity and use in biomonitoring

We compared the use of epilithic, epiphytic, and epipelic diatom communities in stream biomonitoring by investigating species composition and relationships with measured water quality variables in two tributaries of the Grand River, Ontario, Canada. Although ordination showed some separation of the epilithon from other habitats, clear habitat preferences and seasonality were not identified. Canonical correspondence analysis showed that measured water quality variables explained the major gradients in the diatom data for all three habitats. The diatoms from each habitat were strongly related to alkalinity, suspended solids, biological oxygen demand and conductivity, and in addition, epipelic and epiphytic diatoms showed significant relationships with total phosphorus. It was concluded that although some species were more abundant in one habitat than the others, the community structure of the epilithic, epiphytic and epipelic habitats from these streams was not consistently different. The results indicate no apparent benefit to sampling discrete habitats for water quality monitoring using diatoms, however, the best relationship are obtained using the sum of the diatom data from all three habitats.     

INFLUENCE OF CELL VOLUME ON THE GROWTH AND SIZE REDUCTION OF MARINE AND ESTUARINE DIATOMS1

The maximal growth rate (μ_max) of 19 marine and estuarine diatoms decreased with increasing cell volume (V). The relationship between log μ_max (Y) and log V (X) was calculated. Statistical analyses showed that the slope of the equation was not significantly different from those obtained by other researchers and that the 95% confidence intervals of mean μ_max at cell volumes of 10³–10⁵μm³ were not significantly different from those cited in most studies. A new regression line for diatoms was calculated as follows: log μ_max= 0.47–0.14 log V; r =–0.69. The rate of size reduction per generation of the 19 diatom species ranged from 0.03 to 0.87 μm per generation. The rate increased with increasing cell length and cell volume and with decreasing maximum division rate. Statistical analyses showed that the rate was closely related to the cell volume and to the reciprocal of the growth rate. The relationships between maximal growth rate and cell volume and between rate of size reduction and cell volume showed that a diatom with a large volume had a smaller maximal growth rate and a larger rate of size reduction than a diatom with a small volume. The estimates using the equation for the regression line between the rate of size reduction and the reciprocal of maximum division rate indicated that a diatom with a high maximum division rate would need more generation equivalents for a certain size reduction than a diatom with a low maximum division rate, but the periods required for reduction would be approximately equal irrespective of maximum division rate.     

AN INVESTIGATION OF DISTRIBUTIONAL PATTERNS IN THE DIATOM FLORA OF NETARTS BAY,OREGON, BY CORRESPONDENCE ANALYSIS1

Distributional patterns in assemblages of epiphytic and sediment-associated diatoms were investigated in Netarts Bay, Oregon. The method of reciprocal averaging revealed a floristic discontinuity between the epiphytic and sediment samples in ordination space. The basis for this discontinuity was the presence of a large number of sediment-associated taxa that were either very rare or not observed in the epiphytic samples. Within the sediment samples, the diatom flora formed a distributional continuum which had relatively high correlations with mean grain size, a sediment sorting coefficient, and the organic matter content of the sediment. A comparison of the flora in Netarts Bay with floras in other Oregon estuaries indicates that epiphytic, epilithic, and sediment-associated diatom assemblages do not exhibit conspicuous latitudinal changes along the coast of Oregon, and that many of the same taxa can be expected to occur in samples from comparable habitats in estuaries throughout the temperate regions of the world.     

PERIPHYTIC DIATOM ASSEMBLAGES FROM HIGH ARCTIC PONDS1

Epiphytic, epilithic, and surface sediment diatom assemblages were identified and enumerated from 35 study ponds on CapeHerschel (78°37″N, 74°42″W), east-central Ellesmere Island, Canada. All the sites are shallow (maximum depth <2 m), clear, oligotrophic, and freshwater. The ponds freeze completely for 10 months of the year. Major ion concentrations are relatively similar among the 35 sites, although environmental gradients exist. Over 130 diatom taxa from 28 genera were identified in the periphyton samples. Marked differences in species composition were evident among the ponds. Moreover, many of the diatoms exhibited varying degrees of microhabitat specificity. Variance partitioning by canonical correspondence analysis showed that 26% of the total variance exhibited by diatom species composition could be accounted for by the measured environmental variables (i.e. 10.2% by habitat and 15.8% by water chemistry). Pondwater alkalinity best explained the distribution of taxa, and weighted averaging regression and calibration were used to develop a transfer function to infer pondwater alkalinity from the diatom assemblages. Other important environmental variables included [Na⁺] for the epilithic and [SiO₂] for the epiphytic assemblages.     

Epiphytic diatoms in oligotrophc Lake Grane Langsø, Denmark

The epiphytic diatoms on plant-remnants in 10 samples collected in July 1951–1957 from 0.25 m depth in Grane Langsø, Denmark were studied. Averages of their relative frequencies are compared with the relative frequencies of epiphytic diatoms from 11.25 m depth with the aid of a Rank-Abundance diagram. The two graphs were very similar suggesting identity between the epiphytic communities from 0.25 m depth and 11.25 m depth. This suggestion is supported by a highly significant correlation between the relatively abundances of species in the communities from the two depths. Also the populations of the common diatom species Tabellaria flocculosa had the same dimensions of the cells in the two depths and on different host plants, and none of the numerous epiphytic diatoms were ever observed in the plankton of Grane Langsø during five years of regular examination. Overall, the results imply that the littoral plant communities and the pelagic waters offer widely different conditions for growth and survival of diatoms resulting in the development of strictly different communities. The difference between epiphytic and planktonic diatom communities are also likely to become fully expressed due to physical separation of the communities in Grane Langsø resulting from its small size and wind protected location surrounded by forests.     

Developing and Testing Diatom Indicators for Wetlands in the Casco Bay Watershed, Maine, USA

Diatom indicators of wetland condition were developed and tested by assessing human disturbance, water chemistry, and species composition of benthic, epiphytic, and planktonic diatoms from 20 wetlands sampled for 2 years. One sample from each site was randomly selected to form a development data set, while the rest were used as the test data set. Human disturbance indicated substantial differences among wetlands in hydrologic modification, impervious surface, and potential for non-point source contamination. These landscape alterations were related to increases in pH, non-nutrient ions, and nutrients and decreases in dissolved organic carbon and water color. Pre-existing diatom indicators, calculated with autecological information from lakes and aquatic habitats, correlated highly to relevant water chemistry and human disturbance scores. Weighted average models (WAM) of Cl⁻, conductivity, pH, and alkalinity derived with the Maine development data set correlated to relevant water chemistry and human disturbance of the test wetlands. Diatom assemblage attributes that correlated with human disturbance were selected to combine into a multimetric index of biotic condition (IBC). IBCs and WAMs from benthic and epiphytic diatoms were usually more precisely related to relevant environmental factors than planktonic diatoms. These results showed that human disturbance alkalized wetlands, enriched them with nutrients, and diatom assemblages responded to these changes. Indicator development protocols for streams can be readily adapted for use in wetlands.     

Fifteen important questions in the spatial ecology of diatoms

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Growth form analysis of epiphytic diatom communities of Terra Nova Bay (Ross Sea, Antarctica)

Diatoms have been long collected from the Southern Ocean but almost no data exist for epiphytic communities, despite their high ecological significance as an important food source in Antarctic coastal food chains. Here, we present a first growth form analysis of diatoms associated with rhodophyte hosts from Terra Nova Bay, Ross Sea, Antarctica. We performed this study to gather baseline information on the species composition of epiphytic diatom communities, determine the influence of some environmental variables on the diatom distribution patterns, and assess the caveats that must be taken into account in terms of sampling design. Macroalgal material was collected during the Italian Antarctic expeditions between 1990 and 2004. Epiphytic diatoms were studied by means of scanning electron microscopy. In terms of growth forms, there were no significant differences between the diatom communities on the different macroalgal host species. Motile (mainly small-celled Navicula perminuta and other Navicula spp.) and adnate (Cocconeis spp.) diatoms dominated the community throughout the study period. Many of the macroalgal blades examined were also covered by epiphytic animals (calcareous bryozoans, hydroids) over most of their surface, with a significant effect on the associated diatom community structure. Our findings suggest that the bio-physicochemical characteristics of each sampling site affected the epiphytic diatom communities more than the substrate type provided by the macroalgal host or the sampling depth.     

SEASONAL,SUCCESSION OF ALGAE IN RIVERS. I. EXAMPLES FROM THE AVON,A LARGE SLOW-FLOWING RIVER1

The epipelic, epilithic and epiphytic algae in a large slow-flowing river (the Avon) situated in southern England usually exhibited a pronounced bloom during early spring probably in response, to increasing daylength. Although floods caused a reduction in the standing crop of most communities both through scouring action and possibly long-term alteration of microhabitat, they also seemed essential for the renovation of sediment and, rock substrates after periods of rapid algal growth. Because of the attachment characteristics of the predominant epiphytic species, the community attached to Cladophora remained relatively unaffected by flooding. Water velocity may have been important in determining the species composition of some communities, particularly the epiphytic associations. Nutrients probably seldom limited development, but during part of the summer competition with planktonic diatoms for silicon appeared to limit attached algae. During the summer, large scale detachment (which greatly reduced algal density) apparently correlates with a high photosynthetic rate. Algal, numbers in the plankton occasionally decreased in response to a loss of buoyancy of the predominant species.     

Size scaling of extracellular carbonic anhydrase activity in centric marine diatoms

Many microalgae have a surface‐associated extracellular carbonic anhydrase (eCA) that converts HCO₃^? to CO₂ for uptake and subsequent photosynthetic fixation. We investigated eCA activity and assessed its importance for photosynthetic CO₂ supply in six centric diatom species spanning nearly the full range of cell sizes for centric diatoms (equivalent spherical radius 3–67 μm). Since larger cells are more susceptible to diffusion limitation, we hypothesized that eCA activity would increase with cell size as would its importance for CO₂ supply. eCA activity did increase with cell size, increasing with cell radius by a size‐scaling exponent of 2.6 ± 0.3. The rapid increase in eCA activity with cell radius keeps the absolute CO₂ concentration difference between bulk seawater and the cell surface very low (<~0.2 μM) allowing high rates of CO₂ uptake even for large diatoms. Although inhibiting eCA did reduce photosynthesis in the diatoms, there was no overall relationship between the extent of inhibition of photosynthesis and cell size. The only indication that eCA may be more important for larger diatoms was that photosynthesis in the smallest diatoms (<4 μm radius) was only affected by eCA inhibition when CO₂ concentrations were very low, while photosynthesis in some larger diatoms was affected even at typical seawater CO₂ concentrations. eCA is ubiquitous in centric marine diatoms, in contrast to other taxa where its presence is irregularly distributed among different species, and plays an important role in supplying CO₂ for photosynthesis across the size spectrum.     

Gut passage and insect grazer selectivity of lotic diatoms

SUMMARY 1. Grazing experiments were conducted in a small Kentucky stream to determine if epilithic diatom taxa differed in susceptibility to grazing by the larval caddisfly, Neophylax autumnus , and whether ingested taxa varied in digestibility. Live/dead ratios of diatom cells from N. autumnus faecal material were compared with live/dead ratios of cells taken from adjacent epilithic habitats to determine diatom digestibility. Selectivity was studied by comparing relative abundances of diatom taxa on substrata with the relative abundances of these taxa in grazer faecal material.
2. 73% of the diatoms collected from epilithic habitats contained intact chloroplasts and were characterized as live cells. Only 42% of diatoms eliminated in caddisfly faeces were living. The five dominant diatom taxa did not differ in digestibility. Elimination of viable diatoms in grazer faecal material may contribute to diatom drift in streams, and thus supply individuals for downstream recolonization. Failure to document live/dead ratios of diatoms from grazed substrata can lead to overestimation of the digestibility of taxa and misinterpretation of results.
3. Grazers in this investigation were more successful at ingesting large, high-profile diatom taxa (i.e. Cymbella, Meridion and Gomphonema ) and less able to remove small, adnate forms ( Achnanthes minutissima ). Such 'selectivity'may be one mechanism by which A. minutissima maintains dominance in this system.     

Biodiversity of settled material in a sediment trap in the Gulf of Trieste (northern Adriatic Sea)

Phytoplankton succession and sinking rates were studied from January to December 2003 at a coastal station in the Gulf of Trieste (northern Adriatic Sea), 200 m offshore, in a relatively undisturbed area. A conical sediment trap, moored at 15 m depth (water depth 17 m), was used. The hypothesis if the presence of benthic and epiphytic diatoms can lead to an overestimation of the vertical fluxes was tested. To evaluate primary and secondary sedimentation contributions, planktonic, benthic and epiphytic diatoms were distinguished. Benthic species abundance varied throughout the year and it was related to resuspension that strongly influenced sinking rates. All over the year, diatoms were the prevailing class in the trap material accounting for 75.32% of the settled cells, while flagellates represented 24.11%. Dinophyceae and resting cells constituted minor components, accounting for 0.43% and 0.14%, respectively. The gross sedimentation rates ranged from 0.006 × 10⁸ cell m⁻² d⁻¹ in the second week of May to 6.30 × 10⁸ cell m⁻² d⁻¹ in the third week of January with a mean annual value of 1.09 ± 1.43 × 10⁸ cell m⁻² d⁻¹. To the primary sedimentation rate Pseudo-nitzschia seriata of the group “Nitzschia seriata complex” contributed for 49.77% followed by Chaetoceros spp. (23.88%). The major contributor to the secondary sedimentation rate was the diatom Paralia sulcata, accounting for 24.76%. Epiphytic diatoms contributed for 11.19% and 12.27% on annual average gross abundance and biomass, respectively, reaching even 72.04% of gross abundance and 56.06% of gross biomass in the second week of August. The correlation between temperature and the logarithm of the epiphytic biomass was statistically significant, with r = 0.66 and P < 0.001. Both in the cluster analysis and in the PCA four main groups were formed, where benthic and epiphytic species were separately gathered. Planktonic, benthic and epiphytic forms accounted for 50.78%, 36.95% and 12.27%, respectively, calculated on the annual average biomass. Therefore, vertical fluxes can be overestimated of 50% or more if benthic and epiphytic species are not rejected.     

Effects of nutrient (N,P, C) enrichment upon the littoral diatom community of an oligotrophic high-mountain lake

The effects of nutrient additions upon the epilithic diatom communities and the algal standing crop were investigated in the oligotrophic, softwater Lake Piccolo Naret, situated in the Swiss alps. Nutrient-diffusing flower pot substrates were filled with either N (0.15 mol NaNO₃), P (0.015 mol Na₂HPO₄) or C (0.15 mol NaHCO₃) or combinations of them. Twenty-five pots representing eight treatments were placed into the lake in July 1991 and sampled after 42 days of exposure.On the surfaces of all pots containing P we measured higher algal biomasses as on the control pots. The chlorophyll-a maximum of 12.9 µg cm^{– 2} was obtained on NPC pots (0.47 µg cm^{– 2} on control pots). On pots with P, NP or NPC supply high amounts of green algae were detected, also reflected in an increased chl-b/chl-c ratio related to the controls which showed algal communities dominated by diatoms.The diatom communities on the control pots as well as on the pots with N, P and NP had a structure similar to the epilithic community in Lake Piccolo Naret (dominance identity > 58 %). However, the community structures of the diatoms from pots with C addition (C, NC, PC and NPC) differed considerably. This is discussed in view of the cell densities of dominant diatom species. For further comparisons the results of two additional high-mountain lakes are used. By means of a cluster analysis it could be shown that epilithic diatom communities were considerably influenced by C addition, while N and P supply were of minor importance.     

Structure and diversity of intertidal benthic diatom assemblages in contrasting shores: a case study from the Tagus estuary1

The structure of intertidal benthic diatoms assemblages in the Tagus estuary was investigated during a 2‐year survey, carried out in six stations with different sediment texture. Nonparametric multivariate analyses were used to characterize spatial and temporal patterns of the assemblages and to link them to the measured environmental variables. In addition, diversity and other features related to community physiognomy, such as size‐class or life‐form distributions, were used to describe the diatom assemblages. A total of 183 diatom taxa were identified during cell counts and their biovolume was determined. Differences between stations (analysis of similarity (ANOSIM), R = 0.932) were more evident than temporal patterns (R = 0.308) and mud content alone was the environmental variable most correlated to the biotic data (BEST, ρ = 0.863). Mudflat stations were typically colonized by low diversity diatom assemblages (H′ ~ 1.9), mainly composed of medium‐sized motile epipelic species (250–1,000 μm³), that showed species‐specific seasonal blooms (e.g., Navicula gregaria Donkin). Sandy stations had more complex and diverse diatom assemblages (H′ ~ 3.2). They were mostly composed by a large set of minute epipsammic species (<250 μm³) that, generally, did not show temporal patterns. The structure of intertidal diatom assemblages was largely defined by the interplay between epipelon and epipsammon, and its diversity was explained within the framework of the Intermediate Disturbance Hypothesis. However, the spatial distribution of epipelic and epipsammic life‐forms showed that the definition of both functional groups should not be over‐simplified.