Observations on standing crops of epipelic and epipsammic algal communities in Shear Water,Wilts

Methods of estimating the standing crops of epipsammic diatoms have been evaluated and compared with those used for epipelic algal crops.

Good correlations between chlorophyll a content and cell counts were obtained for epipelic populations, but because of the presence of large numbers of decaying cells, inseparable from living cells, such correlation was not obtained with epipsammic algae, even after correction of the pigment estimates for the presence of pheophytin a. The standing crop of epipsammic algae was at all times greater than that of epipelic algae at the same sampling station, and this was attributed to the differential effects of sediment disturbance by burrowing animals and water movements.     

Behavioural versus physiological photoprotection in epipelic and epipsammic benthic diatoms

Benthic diatoms are dominant primary producers in intertidal marine sediments, which are characterized by widely fluctuating and often extreme light conditions. To cope with sudden increases in light intensity, benthic diatoms display both behavioural and physiological photoprotection mechanisms. Behavioural photoprotection is restricted to raphid pennate diatoms, which possess a raphe system that enables motility and hence positioning in sediment light gradients (e.g. via vertical migration into the sediment). The main physiological photoprotection mechanism is to dissipate excess light energy as heat, measured as Non-Photochemical Quenching (NPQ) of chlorophyll fluorescence. A trade-off between vertical migration and physiological photoprotection (NPQ) in benthic diatoms has been hypothesized before, but this has never been formally tested. We exposed five epipelic diatom species (which move in between sediment particles) and four epipsammic diatom species (which live in close association with individual sand grains) to high light conditions, and characterized both NPQ and the relative magnitude of the migratory response to high light. Our results reveal the absence of a significant downward migratory response in an araphid diatom, but also in several raphid epipsammic diatoms, while all epipelic species showed a significant migratory response upon high light exposure. In all epipsammic species the upregulation of NPQ was rapid and pronounced; NPQ relaxation in low light conditions, however, occurred faster in the araphid diatom, compared with the raphid epipsammic species. In contrast, all epipelic species lacked a strong and flexible NPQ response and showed higher susceptibility to photodamage when not able to migrate. While overall our results support the vertical migration-NPQ trade-off, the lack of strong relationships between the capacity for vertical migration and NPQ within the epipsammic and epipelic groups suggests that other factors as well, such as cell size, substrate type and photoacclimation, may influence photoprotective strategies.     

Distribution of benthic diatoms in relation to environmental variables in lowland streams

The composition (% relative abundance) of diatom assemblages from soft bottom sediments was studied at 75 sites situated in 46 rivers, brooks, and ditches in the islands of Hiiumaa and Saaremaa and in the lowland of West Estonia. Although the total number of recorded taxa was 205, the studied diatom assemblages consisted of 54, 55, 48, and 50 constant species in the drainage basin of Moonsund, the Gulf of Riga, Hiiumaa, and Saaremaa, respectively. The habitats of the dominating taxa were heterogenous and the most widespread species were Achnanthidium minutissimum, Martyana martyi, Meridion circulare, Cocconeis placentula, Planothidium lanceolatum, and Amphora pediculus. The Shannon–Weaver diversity (H) index varied from 2.09 to 4.63. Multivariate analyses were used to identify the environmental variables governing the composition and structure of the benthic diatom assemblage. Principal Component Analysis (PCA) and Redundancy Analysis (RDA), based on 56 most abundant taxa, indicated differences in the composition and structure of diatom assemblages between different drainage basins as well as between the upper and lower stream courses. In the headwaters there prevailed small epipsammic diatoms (Martyana, Planothidium, Staurosira, Staurosirella). Different motile epipelic species from the genera Amphora, Navicula, Nitzschia, etc. were distributed abundantly in the lower courses of the streams. There was a positive correlation between order of the stream site and trophic level of water (R=0.35; p<0.05). Along a river system, the increasing order of the stream was accompanied by higher trophic level of water.     

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.     

BENTHIC MARINE DIATOM ASSOCIATIONS: UPPER FLORIDA BAY (FLORIDA) AND ASSOCIATED SOUNDS1

Models of the diatom associations found in upper Florida Bay and adjoining sounds have been constructed utilizing Q modal factor-vector analysis and ecologic diversity indices (Shannon index, number of species, evenness). Four distinct associations were defined using Q-mode factor-vector analysis. Two associations were epiphytic, occurring on Thalassia testudinum König—Association I was characterized by Cocconeis placentula Ehr., Association III by Cylindrotheca closterium (Ehr.) Reim. & Lewin and Cocconeis placentula Ehr. The other two associations were epipelic, occurring on the carbonate mud substratum—Association II was characterized by Cyclotella striata (Kütz.) Grun., Rhopalodia gibberula (Ehr.) O. Müller and Surirella fastuosa (Ehr.) Kütz., Association IV by Fragilaria crotonensis Kitton and Cyclotella striata (Kütz.) Grun. The majority of the 161 species identified were present in both the epiphytic and epipelic assemblages. Only 33 species were restricted to the epiphyton, and 18 species restricted to the epipelon. The epipelic assemblage was significantly more diverse than was the epiphytic assemblage. A general trend of increased diversity away from terrestrial environs toward more open areas of water in both the epipelon and epiphyton was also found.     

SPATIAL AND TEMPORAL VARIATION OF EPIPSAMMIC DIATOMS IN A SPRING-FED BROOK1

Epipsammic diatom communities were sampled and quantified bimonthly from June 1983 to April 1985 at four sites in a small brook in northwest Ohio. The primary objective was to determine the extent of temporal and spatial variability in the epipsammon. The four sites, approximately 250 m apart, differed in current velocity, illumination and the amount of sand-associated detritus. Diatoms were significantly more abundant in the portion of the brook flowing through an unshaded marsh than at three heavily shaded sites located upstream. Fragilaria leptostauron (Ehr.) Hust. and Achnanthes lanceolata var. dubia Grun. dominated the epipsammic assemblage at all sites throughout the entire study period. Only Meridion circulare (Grev.) Ehr. displayed a marked seasonal distribution. Although the same species were generally found at all sites, there were sufficient differences in relative abundance that communities could be discriminated according to site. Using two canonical variate axes, all 48 samples were correctly assigned to their proper sampling site based on community composition. Data from this epipsammic assemblage support the idea developed from the river continuum concept that species comprising riverine benthic assemblages continually persist and rarely become completely absent.     

Influence of the colonizing substrate on diatom assemblages and implications for bioassessment: a mesocosm experiment

Although diatoms are important bioindicators of ecological quality, their ecological traits are still not well understood. A major issue is that of substrate preferences, which may result in differences in production, and assemblage structure and composition, and which should therefore be taken into account for ecological quality assessment studies. Thus, in this work, the periphyton grown on sand and ceramic tiles in indoor controlled channels were compared to understand whether substrate differences lead to differences in: periphyton production (chlorophyll-a), chlorophyll-b and c concentrations, diatom assemblages (diversity-Shannon-Wiener, cell density, taxonomic composition, trait proportions), and ecological quality assessments (IPS-‘Indice de Polluosensibilité Spécifique’). A combined inoculum of periphyton from four Portuguese streams was introduced to the running channels (six sand and six tile) and left to colonize for 35 days. Epilithic (tiles) and epipsammic (sand) assemblages were sampled at days 14 and 35. We verified that there were no differences in chlorophyll-a concentration over time and between substrates. On both sampling occasions, the epipsammic assemblages had higher concentration of chlorophyll-c and diatom density but without significant differences over time in each substrate. The taxonomic composition was different between substrates and over time. However, these differences were not reflected in ecological quality assessment. The diversity was also similar between substrates in both sampling occasions, but it was higher at day 14. Mobile and stalked species were more abundant over the entire study and differed significantly between substrates, with the epipsammic assemblages presenting higher abundances of both traits. We concluded that the colonizing substrate influences diatom assemblages but not the ecological quality assessment.     

Early-20th Century Environmental Changes Inferred Using Subfossil Diatoms from a Small Pond on Melville Island, N.W.T., Canadian High Arctic

Diatom-based paleolimnological studies are being increasingly used to track long-term environmental change in arctic regions. Little is known, however, about the direction and nature of such environmental changes in the western Canadian high Arctic. In this study, shifts in diatom assemblages preserved in a ²¹⁰Pb-dated sediment core collected from a small pond on Melville Island, N.W.T., were interpreted to record marked environmental changes that had taken place since the early 20th century. For most of the history of the pond recorded in this core, the diatom assemblage remained relatively stable and was dominated by Fragilaria capucina. A major shift in species composition began in the early-20th century, with a sharp decline in F. capucina and a concurrent increase in Achnanthes minutissima. In the last 20 years, further changes in the diatom assemblage occurred, with a notable increase in the Nitzschia perminuta complex. The assemblage shifts recorded at this site appear to be consistent with environmental changes triggered by recent climatic warming.     

Distribution of epipelic diatoms in artificial fishponds along environmental and spatial gradients

Although epipelic diatoms play a key role in primary production of many ecosystems, many aspects of their biodiversity, ecology and geographical distribution are poorly understood. The present study is based on sampling of 45 man-made fishponds in the Czech Republic covering an environmental gradient from oligo/dystrophic highland ponds within protected areas to the eutrophic/hypertrophic lowland ponds used for intensive fish production. Diatom distribution patterns assessed using biomass and species composition variables were assessed along environmental and geographical gradients. In total, 185 epipelic diatom taxa were found in the investigated samples. The differences in species composition between sites were correlated with environmental parameters, but not with the geographic distance of the localities. This pattern might suggest that niche-based control, rather than the effect of dispersal limitation, is the main driving force in the species composition of epipelic diatoms in fishponds. The alpha-diversity of sites correlated with altitude, nitrogen and chlorophyll a concentrations but did not correlate with the area of the ponds. The significant relationships between local abundance of species and their regional occupancy were very similar to previous studies of diatoms in boreal streams. In addition, these data concur with patterns known for multicellular organisms suggesting that in this respect diatoms may not differ from groups of organisms with larger body sizes. Handling editor: J. Padisak     

Microscale imaging sheds light on species-specific strategies for photo-regulation and photo-acclimation of microphytobenthic diatoms

Intertidal microphytobenthic (MPB) biofilms are key sites for coastal primary production, predominantly by pennate diatoms exhibiting photo-regulation via non-photochemical quenching (NPQ) and vertical migration. Movement is the main photo-regulation mechanism of motile (epipelic) diatoms and because they can move from light, they show low-light acclimation features such as low NPQ levels, as compared to non-motile (epipsammic) forms. However, most comparisons of MPB species-specific photo-regulation have used low light acclimated monocultures, not mimicking environmental conditions. Here we used variable chlorophyll fluorescence imaging, fluorescent labelling in sediment cores and scanning electron microscopy to compare the movement and NPQ responses to light of four epipelic diatom species from a natural MPB biofilm. The diatoms exhibited different species-specific photo-regulation features and a large NPQ range, exceeding that reported for epipsammic diatoms. This could allow epipelic species to coexist in compacted light niches of MPB communities. We show that diatom cell orientation within MPB can be modulated by light, where diatoms oriented themselves more perpendicular to the sediment surface under high light vs. more parallel under low light, demonstrating behavioural, photo-regulatory response by varying their light absorption cross-section. This highlights the importance of considering species-specific responses and understanding cell orientation and photo-behaviour in MPB research.     

Hydrological controls on the diatom assemblage of a seasonal arctic river: Boothia Peninsula,Nunavut, Canada

This study characterizes the melt season diatom assemblage of a middle arctic river with respect to hydrological conditions. In addition, the potential to identify species that show a strong affinity for the lotic environment provides an opportunity to interpret stratigraphic changes in these species in the lake sedimentary record in terms of past hydrological change. Understanding long-term hydrological variability is critical for assessing both current and future environmental change. Significantly higher relative abundances of Achnanthes minutissimaKützing, Fragilaria capucina var. vaucheriae (Kützing) Lange-Bertalot, Diatoma tenuisAgardh, Cymbella arctica(Lagerstedt) Schmidt, C. minutaHilse ex. Rabenhorst, C. silesiaca Bleisch and Encyonema fogedii Krammer in the lotic environment throughout the 2001 growing season compared to the lacustrine sedimentary record suggest that these species characterize the Lord Lindsay River diatom assemblage. Comparison of seasonal abundances of these taxa to hydrological parameters including discharge, electrical conductivity, and water temperature reveal key information about the character of this community. The fact that the river diatom assemblage changes very little throughout the sampling period, despite major changes in hydrological conditions, suggests a degree of resilience and inherent structure in the community. However, a decrease in diatom biomass in response to rapid and dramatic changes in hydrological conditions following a major rainfall event suggests that a threshold tolerance may exist, with potentially important implications for interpreting stratigraphic changes in the paleoenvironmental record.     

The quantitative variability and monosaccharide composition of sediment carbohydrates associated with intertidal diatom assemblages

The extracellular secretions of epipelic diatoms (Bacillariophyceae) axe an important source of carbohydrates on intertidal sediments. For analytical purposes, sediment carbohydrates have been operationally separated into colloidal and bulk fractions that are often assumed to be similar in their chemical properties. However, there has been little investigation into the nature of the two fractions. In this study, carbohydrate fractions were sampledin situ, isolated, purified and biochemically characterised using gas chromatography-mass spectrometry (GC-MS). Both carbohydrate fractions were found to contain similar sugars although in different proportions. Glucose represented more than 80% of the monosaccharides identified in the colloidal carbohydrate fraction while only 37% of monosaccharides present in the bulk carbohydrate fraction. Colloidal carbohydrate concentrations showed short-term variability and were correlated with diatom biomass (as chlorophylla) suggesting the colloidal fraction is labile and may be of recent origin, perhaps representing diatom activity. Concentrations of the bulk carbohydrate fraction did not show significant short-term variation and was therefore more refractory. This combination of biochemical and field data suggested that the bulk and colloidal carbohydrate fractions were chemically and physically different. These findings have relevance to studies of estuarine carbon cycling.Abbreviations LTSEM Low-temperature scanning electron microscopy - GC-MS Gas chromatography-mass spectroscopy - EPS Extracellular polymeric substances     

Diatom ecological preferences in a shallow temperate estuary (Ria de Aveiro,Western Portugal)

The study of the diatom ecological preferences was conducted from January 2002 to June 2003 in Canal de Mira, Ria de Aveiro, Western Portugal. Three sampling stations along a salinity gradient were sampled monthly, in new moon, at high and low tide. Salinity, temperature, pH, dissolved oxygen and nutrient contents were measured for each sampling station; chlorophyll a and diatom diversity and abundance were also evaluated. Canonical correspondence analysis was used to identify the environmental variables governing the composition and structure of diatom assemblage. The variation in the species data among the different reaches was strongly determined by the salinity spatial gradient and by the temperature temporal gradient. The lower reaches were dominated by marine species (e.g. Auliscus sculptus, Chaetoceros densus, Fallacia forcipata,Licmophora flabellata, L. grandis, Surirella comis), while in the most upstream station typical freshwater species dominated (e.g. Caloneis permagna, Cymatopleura solea, Cymbella tumida, Gomphonema longiceps, Pinnularia stommatophora, Stauroneis smithii). Weighted averaging was used to estimate optima and tolerances of some diatom taxa for the most influential variables. It was possible to establish groups oftaxa with defined and distinctive salinity and temperature preferences.     

Predicting Epipelic Diatom Exopolymer Concentrations in Intertidal Sediments from Sediment Chlorophyll a

Abstract In many intertidal cohesive—sediment habitats, epipelic diatoms are the dominant microphytobenthic organisms. In such sediments, concentrations of colloidal carbohydrate [including the exopolymeric substances (EPS) produced by diatoms during motility] are closely correlated with the biomass (chlorophyll a) of epipelic diatoms. A model describing this relationship (log (conc. coll. carbo. + 1) = 1.40 + 1.02(log (chl. a conc. + 1)) was derived from published data. It was validated against published and unpublished data from 6 different estuaries, and accounted for 64.6% of the variation in sediment colloidal carbohydrate concentrations. The model was valid for intertidal habitats with cohesive sediments where epipelic diatoms constituted >50% of the microphytobenthic assemblage. In sites with noncohesive sediments, or where the microphytobenthic assemblage was dominated by other algal groups, the model was not applicable. The mean percentage of EPS in colloidal carbohydrate extracts varied between 11 and 37% for axenic cultures of epipelic diatoms (with higher values obtained during stationary phase), and between 22.7% and 24.3% for natural sediments dominated by epipelic diatoms. Assuming an EPS percentage of 25% in colloidal extracts yielded an EPS chl. a ratio of 2.62:1. Maximum rates of EPS production in diatom cultures occurred at the beginning of stationary phase (1.6–5.09 μg EPS μg⁻¹ chl a d⁻¹), with Nitzschia sigma having a significantly (P < 0.05) higher rate of production than N. frustulum, Navicula perminuta and Surirella ovata. Similar rates of EPS production were measured in the field. The dynamics of EPS production and loss on mudflats is discussed, with reference to the model and these production rates. Received: 25 February 1997; Accepted: 23 May 1997     

Characterization of the diatom flora in the Lower Mountain Fork (Oklahoma,USA), a novel regulated river with a disjunct population of the diatom Didymosphenia geminata (Bacillariophyta)

Regulated rivers are novel ecosystems with altered temperature and flow regimes that can be used to test distribution patterns of microscopic organisms, such as diatoms. Our objective was to describe the spatial and seasonal patterns of diatoms in a cold-water, oligotrophic river within a region of warm-water, mesotrophic rivers. The Lower Mountain Fork, in south-east Oklahoma (USA), is maintained as a year-around, stocked fishery by the release of cold, hypolimnetic water from Broken Bow Lake and is the southern-most known site of Didymosphenia geminata in North America. Epilithic diatoms were sampled six times at nine sites over a distance of 15.5 km and, within this area, 27 times at the site of the main Didymosphenia bloom. Percentage composition data were analysed for assemblage composition using multivariate analysis, nutrient specificity using a diatom-based metric, and species associations using similarity profiles. Eighty-eight taxa were found, of which 10 were unidentifiable and included local undescribed species and species clusters. Three species [Gomphonema (parvulum morph), Achnanthidium rivulare and Achnanthidium minutissimum] comprised over 60% of the diatom abundance at all sites, and downstream and seasonal patterns were evident for both these and less abundant taxa. Notably, diatom assemblages in the three sites below the dam were similar to that at the lowermost site, below a much smaller dam. The oligotraphentic diatom assemblage reflected the water chemistry of the river. The Didymosphenia bloom had been scoured by a large spate prior to the study and the species was present at two of the nine sites in low numbers but failed to bloom during the study, possibly because of a trend towards increasing phosphorus concentrations in the reservoir (Didymosphenia blooms under low P concentrations). No other species shared Didymosphenia’s distribution pattern over the study reach, highlighting the novelty of Didymosphenia’s presence in the river.     

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.     

Psammodiscus nov. gen. Based on Coscinodiscus nitidus

The epipsammic diatom Coscinodiscus nitidus differs from typicalCoscinodiscus species in areola arrangement and structure, closingmembrane structure, rimoportula arrangement, and habitat. Severalsimilarities exist between it and Rhaphoneis species but, whilethe relationship between these is close, the two cannot be consideredcongeneric. A new genus, Psammodiscus, is created to includeC. nitidus. Coscinodiscus nitidus, Rhaphoneis, Psammodiscus nov. gen., epipsammic diatom, diatom taxonomy     

EPIPELIC AND EPILITHIC DIATOM ASSEMBLAGES IN MONTEZUMA WELL NATIONAL MONUMENT,ARIZONA1

Epipelic and epilithic diatom assemblages in Montezuma Well National Monument, Arizona, were identified and analyzed by H' (species diversity), species number, J' (evenness), relative abundance, SIMI (similarity index), and DI (dissimilarity index) values. The epipelic assemblage was found to be a tri-dominant association of Cyclotella pseudostelligera f. parva Czar. & Blinn, Cocconeis placentula var. lineata (Ehr.) V.H. and Achnanthes linearis (W. Sm.) Grun. The epilithic assemblage was a tri-dominant association of Rhoicosphenia curvata (Kütz.) Grun. ex Rabh., Cy. Pseudostelligera f. parva and Synedra ulna (Nitz.) Ehr. Both associations persisted throughout the year. Of the 83 taxa observed, 51 were common to both assemblages, 24 were restricted to the epipelon, and 8 were restricted to the epilithon. On a yearly basis, the epipelic assemblage exhibited less diversity (H') than the epilithon, even though the latter exhibited fewer taxa. The use of mean diversity values was also determined to be unacceptable. Cy. Pseudostelligera f. parva and S. ulna appeared more abundant under high light intensity and duration, Fragilaria construens var. venter (Ehr.) Grun. favored low light intensity and duration, whereas A. linearis, R. curvata and Co. placentula var. lineata were tolerant of a variety of intensities and durations. On a yearly basis, both assemblages exhibited slight similarity (SIMI, DI). The use of SIMI values for comparing assemblages with few highly abundant taxa was determined to be unacceptable and under these conditions, the use of DI was recommended.     

Flood disturbance effects on benthic diatom assemblage structure in a semiarid river network

Disturbances such as floods and droughts play a central role in determining the structure of riverine benthic biological assemblages. Extreme disturbances from flash floods are often restricted to part of the river network and the magnitude of the flood disturbance may lessen as floods propagate downstream. The present study aimed to characterize the impact of summer monsoonal floods on the resistance and resilience of the benthic diatom assemblage structure in nine river reaches of increasing drainage size within the Gila River in the southwestern United States. Monsoonal floods had a profound effect on the diatom assemblage in the Gila River, but the effects were not related to drainage size except for the response of algal biomass. During monsoons, algal biomass was effectively reduced in smaller and larger systems, but minor changes were observed in medium systems. Resistance and resilience of the diatom assemblage to floods were related to specific species traits, mainly to growth forms. Tightly adhered, adnate and prostrate species (Achnanthidium spp., Cocconeis spp.) exhibited high resistance to repeated scour disturbance. Loosely attached diatoms, such as Nitzschia spp. and Navicula spp., were most susceptible to drift and scour. However, recovery of the diatom assemblage was very quick indicating a high resilience, especially in terms of biomass and diversity. Regional hydroclimatic models predict greater precipitation variability, which will select for diatoms resilient to bed‐mobilizing disturbances. The results of this study may help anticipate future benthic diatom assemblage patterns in the southwestern United States resulting from a more variable climate.