DISTRIBUTION OF INTERTIDAL DIATOMS ASSOCIATED WITH SEDIMENTS IN YAQUINA ESTUARY,OREGON1,2

Sediment samples and environmental data were collected from November 1973 to August 1974 to analyze the distribution of sediment-associated diatom assemblages relative to vertical, horizontal, and seasonal gradients in Yaquina Estuary, Oregon. The distribution of diatoms was regulated primarily by mean salinity and characteristics of the sediment, i.e., mean sediment size and percentage of organic carbon. Prominent taxa in the river above Yaquina Bay exhibited overlapping distributions along the salinity gradient to a location in brackish water where the mean salinity was approximately 5%_o. At this salinity, a relatively sharp discontinuity in the diatom flora existed which appeared to represent the biochemical and biophysical mechanisms involved in the osmotic regulation of fresh- and brackish-water diatoms. Relatively large disparities m the structure of diatom assemblages were found within relatively small areas of Yaquina Bay. These differences were attributed to properties of the sediment. Differences in diatom assemblages relative to variations in light intensity, water temperature and exposure to intertidal emergence were not apparent from the analysis of field data.     

PATTERNS OF DISTRIBUTION OF DIATOM ASSEMBLAGES ALONG ENVIRONMENTAL GRADIENTS IN THE SEVERN RIVER ESTUARY,CHESAPEAKE BAY,MARYLAND1

Distribution patterns of diatoms from natural substrates and surface waters of the Severn River, Maryland, are related to salinity and seasonal gradients through time and spare by minimum spanning tree cluster analysis and principal components analysis. Assemblages were found to be distributed in a complex but orderly fashion, with primary discontinuities occurring between seasons, and between planktonic and attached forms. Within seasons, benthic groupings are distributed as continua along a horizontal salinity gradient, with one discontinuity at a location with a mean salinity of approximately 7‰, Downstream from this discontinuity, most species do not follow the changing salinity patterns from season to season, and are considered euryhaline for a salinity range of 8–16‰. Upstream from this discontinuity there is a relatively distinct and persistent assemblage of oligohaline indicators. Evidence derived by comparing this study with similar studies from Oregon estuaries indicates that similar distribution patterns, community structures and taxa can be expected to occur in estuaries on the east and west coasts of North America.     

PATTERNS OF DISTRIBUTION OF DIATOM ASSEMBLAGES ALONG ENVIRONMENTAL GRADIENTS IN THE SEVERN RIVER ESTUARY,CHESAPEAKE BAY,MARYLAND1

Distribution patterns of diatoms from natural substrates and surface waters of the Severn River, Maryland, are related to salinity and seasonal gradients through time and space by minimum spanning tree cluster analysis and principal components analysis. Assemblages were found to be distributed in a complex but orderly fashion, with primary discontinuties occurring between seasons, and between planktonic and attached forms. Within seasons, benthic groupings are distributed as continua along a horizontal salinity gradient, with one discontinuity at a location with a mean salinity of approximately 7%o. Downstream from this discontinuity, most species do not follow the changing salinity patterns from season to season, and are considered euryhaline for a salinity range of 8–16%o. Upstream from this discontinuity there is a relatively distinct and persistent assemblage of oligohaline indicators. Evidence derived by comparing this study with similar studies from Oregon estuaries indicates that similar distribution patterns, community structures and taxa can be expected to occur in estuaries on the east and west coasts of North America.     

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.     

DISTRIBUTION OF EDAPHIC DIATOMS IN A MISSISSIPPI SALT MARSH: A CANONICAL CORRELATION ANALYSIS1

A canonical correlation analysis was performed to examine the degree to which edaphic diatom distribution in Graveline Bay Marsh, Mississippi could be related to environmental factors. It was possible to collapse the complex data base into two interpretable, orthogonal dimensions and to identify five distinct diatom communities. Diatom distribution on the marsh was primarily regulated by elevation and height of the overstory spermatophyte canopy. Although redundancy in the species data given the environmental data was relatively low (23% for the first two canonical variables), similar values were obtained for studies of benthic and planktonic diatoms in Yaquina Estuary, Oregon. The present study represents the first application of multivariate statistics to the study of salt marsh diatoms and demonstrates its usefulness in an environment where gradients are difficult to perceive.     

STUDIES ON MARINE PLANKTON DIATOMS. II. RESTING SPORE MORPHOLOGY1

A morphological study of resting spores in five marine planktonic diatom species using electron microscopy indicates that Bacteriastrum delicatulum Cleve and Leptocylindrus danicus Cleve spores bear little resemblance, to their vegetative cells. Detonula confervacea (Cleve) Gran and Stephanopyxis turris (Grev. & Arn.) Ralfs spores have several features in common with their vegetative cells, and Rhizosolenia setigera Brightwell lies between the two extremes. The function of resting spores in relation to diatom life cycles is briefly discussed. Spore formation may be a primitive characteristic in the life cycle and may no longer have significant survival value for the species.     

DIATOM ASSOCIATIONS IN YAQUINA ESTUARY,OREGON: A MULTIVARIATE ANALYSIS1

A cluster analysis and a principal components analysis were performed to examine the degree to which the diatom flora of Yaquina estuary can be partitioned into discrete associations and to relate the composition of the flora to selected physical properties of the estuary. The species composition of the diatom flora during the winter was more closely associated with salinity and intertidal exposure them in the summer. A cluster of freshwater taxa obtained for the winter data was the most conspicuous diatom association defined by the cluster analysis. Although taxa within a cluster had more similar distributions than taxa from different clusters, there was considerable variation in the distributions of marine and brackish water taxa within a multispecies cluster.     

PRODUCTION OF EPIBENTHIC DIATOMS IN A SOUTHERN CALLIPORNIA IMPOUNDED ESTUARY1, 2

In situ epibenthic diatom productio was determined for four sites within Bolsa Bay (Orange Coutry), California, to assess the effect of estuative impoundment by a tidal gate. Mean annual values of 26 and 41 mg C m^?2 h^?1 were found in the finner sediment of the outer bay whereas mean annual values of 41 and 59 mg C m^?2 h ^?1 were found in the coarser, more stable sediment of the inner bay. Low production in the vicinity of the mouth of a flood control channel is thought to be caused by heavy metal inhibition in conjuction with sediment instability, a shl/allow photic zone and heavy grazing pressure. Seasonal production peaks in spring and falll were found to be closely correlated with the concentration of nitrate and silicate but not with phosphate. This study shows that even though Bolsa Bay has been cut off from the sea, drained and subjected to fill, it is as productive as other coastal wetlands throughout the world.     

THE ECOLOGICAL IMPLICATIONS OF GROWTH FORMS IN EPIBENTHIC DIATOMS 1

This paper evaluates the utilisation of space by epibenthic diatom cells, as a response to environmental variations. The aggregation pattern of five species of epibenthic diatoms was quantified and compared to provide evidence for the significance of cell motility as an adaptive mechanism for space occupation and monopoly. The epibenthic diatoms included (1) non-mobile colonial species forming either fan-shaped (Synedra tabulata (Ag)Kz.) or arborescent (Gomphonema kamtschaticum var. californicum Grun.) colonies; (2) slow-moving (Cocconeis costata Greg, and Amphora pusio Cl.), and (3) fast-moving (Navicula direct a (W. Sm.) Ra.) non-colonial species. The aggregation pattern of S. tabulata did not vary significantly among six different light intensities manipulated in nature. The major patterns of aggregation were identified using analysis of covariance and dummy-variable regression. Highly mobile N. directa are significantly less aggregated than the four other diatom species. Non-mobile and slow-moving species show a similar, highly aggregated pattern. The occurrence of two patterns of spatial dispersion indicates that growth forms bear far-ranging ecological implications with respect to colonization strategies, immigration, and possibly impact by grazers. An integrated model of growth form characteristics, biological properties, and ecological implications is presented for epibenthic diatoms.     

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.     

INTRASPECIFIC COMPARISONS OF NITRATE UPTAKE IN THREE MARINE DIATOMS1

Short-term (within 5 min) and long-term (≤2 h) rates of nitrate uptake were determined for the marine diatoms, Nitschiella longissima (Cleve), Skeletonema costatum (Greville) Cleve and Asterionella japonica (Cleve). Pigment levels, cell carbon, nitrogen and cell volume were also determined for cells in the logarithmic and stationary phases of growth. For each species, one clone isolated from oligotrophic coastal water and one clone isolated from eutrophic coastal water were compared. Long-term NO₃^? uptake typically followed saturation kinetics describable by the Michaelis-Menten expression. Under experimental conditions, half-saturation constants ranged from 0.6 to 2.2 μM NO₃^?. Generally, the oligotrophic clones had lower Ks and V_max (on a per cell basis) than their eutrophic counterparts, though this was only statistically significant in one pair of clones. Eutrophic and oligotrophic clones also differed in their short-term response to nutrient addition; oligotrophic clones showed greatest rate of uptake at the lowest nitrate addition while uptake by eutrophic clones increased with increasing nitrate concentration. However, all clones had very similar V_max values expressed on a dry weight basis. Under N-starvation, cellular C and pigment levels (and N to a lesser extent) generally declined more in eutrophic than in oligotrophic clones. While the differences between inshore and offshore clones were not great, the results are consistent with the hypothesis that eutrophic waters support algae which grow faster and are less conservative biochemically than cells in oligotrophic waters.     

EXTREMELY HIGH GROWTH RATE OF THE SMALL DIATOM CHAETOCEROS SALSUGINEUM ISOLATED FROM AN ESTUARY IN THE EASTERN SETO INLAND SEA,JAPAN1

Small single‐celled Chaetoceros sp. are often widely distributed, but frequently overlooked. An estuarine diatom with an extremely high growth potential under optimal conditions was isolated from the Shinkawa‐Kasugagawa estuary in the eastern part of the Seto Inland Sea, western Japan. It was identified as Chaetoceros salsugineum based on morphological observations. This strain had a specific growth rate of 0.54 h^?1 at 30°C under 700 μmol · m^?2 · s^?1 (about 30% of natural maximal summer light) with a 14:10 L:D cycle; there was little growth in the dark. However, under continuous light it grew at only 0.35 h^?1 or a daily specific growth rate of 8.4 d^?1. In addition, cell density, chlorophyll a, and particulate organic carbon concentrations increased by about 1000 times in 24 h at 30°C under 700 μmol · m^?2 · s^?1 with a 14:10 L:D cycle, showing a growth rate of close to 7 d^?1. This very rapid growth rate may be the result of adaptation to this estuarine environment with high light and temperature. Thus, C. salsugineum can be an important primary producer in this estuary in summer and also an important organism for further physiological and genetic research.     

STUDIES ON MARINE PLANKTON DIATOMS. III. STRUCTURE AND CLASSIFICATION OF GOSSLERIELLA TROPICA1

The structure of the tropical oceanic diatom Gossleriella tropica Schütt was examined in the light and electron microscopes. The species is characterized by a primarily poroid valve, surface, it single central labiate process and a unique ring of spines attached to the cingulum. Placement of the genus in the family Rhizosoleniaceae is suggested. Although G. tropica is widely distributed in the world's oceans, it is usually confined to the lower photic zone, salinities of 34.5–35.5% and temperatures of 15–30 C.     

THE RELATIVE PHOTOSYNTHETIC EFFICIENCY OF ASTERIONELLA FORMOSA HASS. (BACILLARIOPHYTA) IN NATURAL PLANKTON ASSEMBLAGES1

The relative photosynthetic efficiencies of net vs. nanoplankton and diatom vs. non-diatom plankton were estimated weekly in Woodcock Creek Reservoir, Pennsylvania, from April to November 1979, using carbon-14 in conjunction with size fractionation and germanium as a diatom inhibitor. The nanoplankton contributed a proportionally greater share of the community carbon assimilation than their share of the community biomass would indicate. Diatoms, specifically Asterionella formosa Hass. were photosynthetically inefficient and contributed significantly less than expected. The mechanism of periodic diatom dominance in lakes remains unexplained.     

BLOOMS OF SURF-ZONE DIATOMS ALONG THE COAST OF THE OLYMPIC PENINSULA,WASHINGTON. VI. DAILY PERIODICITY PHENOMENA ASSOCIATED WITH CHAETOCEROS ARMATUM IN ITS NATURAL HABITAT1,2

Microscope observations of chains of Chaetoceros T. armatum T. West, collected, from its natural habitat, at Copalis Beach, Washington, have been made. A regular daily periodicity in the occurrence of cell division stages and length of the chains was found on each occasion (November, February, April, May, August). During the night, short chains of 3 cells predominated. In early morning hours, average chain lengths increased due to cell elongation, and nuclear and protoplast division occurred. This correlated with the time of day when cell masses of C. armatum first appeared at the surface of the water. In the late afternoon, the new silica valves of the daughter protoplasts were formed. Formation of special bristle-bearing end valves effected the separation of long chains into shorter chains composed predominantly of 3 cells. (Cultured material was used to verify some of these stages.) This correlated with the time of day when cell masses disappeared from the water surface and dispersed.     

THE EVOLUTION OF ELONGATE SHAPE IN DIATOMS1

Diatoms have been classified historically as either centric or pennate based on a number of features, cell outline foremost among them. The consensus among nearly every estimate of the diatom phylogeny is that the traditional pennate diatoms (Pennales) constitute a well‐supported clade, whereas centric diatoms do not. The problem with the centric–pennate classification was highlighted by some recent analyses concerning the phylogenetic position of Toxarium, whereby it was concluded that this “centric” diatom independently evolved several pennate‐like characters including an elongate, pennate‐like cell outline. We performed several phylogenetic analyses to test the hypothesis that Toxarium evolved its elongate shape independently from Pennales. First, we reanalyzed the original data set used to infer the phylogenetic position of Toxarium and found that a more thorough heuristic search was necessary to find the optimal tree. Second, we aligned 181 diatom and eight outgroup SSU rDNA sequences to maximize the juxtapositioning of similar primary and secondary structure of the 18S rRNA molecule over a much broader sampling of diatoms. We then performed a number of phylogenetic analyses purposely based on disparate sets of assumptions and found that none of these analyses supported the conclusion that Toxarium acquired its pennate‐like outline independently from Pennales. Our results suggest that elongate outline is congruent with SSU rDNA data and may be synapomorphic for a larger, more inclusive clade than the traditional Pennales.     

THE DISTRIBUTION OF EDAPHIC DIATOMS ALONG ENVIRONMENTAL GRADIENTS OF A LOUISIANA SALT MARSH1

Edaphic (sediment-associated) diatoms were collected from five sites along a gradient from slightly brackish to saline marsh on 30 November 1977 and 29 April 1978. Several environmental factors change along this gradient in addition to salinity, including tidal flushing and percent organic content of the soil. A total of 112 taxa were identified, with Amphora exigua Greg., Navicula phyllepta Kütz., N. salinarum Grun., and N. tripunctata (Müll.) Bory being among the five most abundant taxa on both sampling dates. The taxonomic composition of the diatom flora was very similar to those observed for other salt marshes located along the Gulf and Atlantic coasts. Diatom taxa were generally euryhaline and distributed continuously along the gradient from brackish to saline marsh. Species diversity (H¹) and the number of taxa in a sample (S) were also similar to those reported for other North American salt marshes. The number of taxa in a sample was higher in the saline than in the brackish area on both sampling dates while species diversity was greater in the saline area only in April.     

EFFECTS OF LIGHT AND SUBSTRATE ON THE BENTHIC DIATOMS IN AN OLIGOTROPHIC LAKE: A COMPARISON BETWEEN NATURAL AND ARTIFICIAL SUBSTRATES1

Benthic diatoms form a particularly important community in oligotrophic lakes, but factors influencing their distribution are not well known. This study reports the depth distribution of living motile and total diatoms (living plus dead diatoms) on both natural (from sand to fine organic mud) and artificial substrates in an oligotrophic lake. On artificial substrates, motile diatom densities peaked in abundance (24–30 cells · mm^?2) between 0.6 and 1.9 m depth; on natural sediment surfaces, motile diatoms were generally more numerous and peaked in abundance (925 cells · mm^?2) at 1.3 m depth. Total diatom densities on artificial substrates were highest (1260 valves · mm^?2) at 0.6 m depth, with very low values below 3 m depth; on natural sediment surfaces, total diatom abundances were generally much higher (21600 valves · mm^?2) at 3 m depth and declined gradually with depth. Significant relationships were found between light and diatom densities on the artificial substrate. Ordination analysis indicated that substrate type significantly correlated with the variation of diatom composition on artificial and natural substrates. Our results suggest that in oligotrophic lakes, light influences benthic diatom abundance, whereas substrate type has more influence on benthic diatom composition.     

STUDIES IN PENNATE DIATOMS: VALVE MORPHOLOGIES OF LICMOPHORA AND CAMPYLOSTYLUS1

Using scanning electron and light, microscopy, several hundred specimens of Campylostylus normanianus (Grev.) Gerloff, Licmophora abbreviata Agardh, L. gracilis var. anglica (Kütz) Per. et Per., and L. flabellata (Carm.) Agardh were examined to elucidate their valve morphology. These species were found to be heterovalvar with respect to the presence of the labiate process in the basal apices of the cell, although one was always present at the head pole of each valve. This form of heterovalvar may be one basis for partitioning the large and variable family Diatomaceae. Because of the similar valve morphologies exhibited by C. normanianus and Licmophora species examined, it is recommended that Campylostylus normanianus be placed in the older genus Licmophora, as L. normaniana (Grev.) Wahrer.