A coded checklist and ecological indicator values of freshwater diatoms from The Netherlands

This first comprehensive checklist of the diatoms from fresh and weakly brackish water in The Netherlands comprises 948 taxa, belonging to 776 species in 56 genera. The generaNavicula, which has a very wide ecological amplitude, andNitzschia, which has many pollution tolerant species, are most numerous. Each taxon is identified with a unique eight-letter code, to facilitate computer processing of data. Ecological indicator values for pH, salinity, nitrogen uptake metabolism, oxygen, saprobity, trophic state and moisture are presented.     

LINKING BENTHIC ALGAL BIOMASS TO STREAM SUBSTRATUM TOPOGRAPHY1

The physical properties of substrata significantly influence benthic algal development. We explored the relationships among substratum surface texture and orientation with epilithic microphytobenthic biomass accumulation at the whole‐substratum and micrometer scales. Unglazed clay tiles set at three orientations (horizontal, vertical, and 45°), and six substrata of varying surface roughness were deployed in a prairie stream for 3 weeks. Substrata were analyzed for loosely attached, adnate, and total benthic algal biomass as chl a, and confocal laser scanning microscopy was used to measure substrata microtopography (i.e., roughness, microscale slope angles, and three‐dimensional surface area). At the whole‐substratum level, vertical substrata collected significantly (P < 0.05) less algal biomass, averaging 34% and 36% less than horizontal and 45° substrata, respectively. Benthic algal biomass was also significantly less on smoother surfaces; glass averaged 29% less biomass than stream rocks. At the microscale level, benthic algal biomass was the greatest at intermediate values, peaking at a mean roughness of approximately 17 μm, a mean microscale slope of 50°, and a projected/areal surface area ratio of 2:1. The proportion of adnate algae increased with surface roughness (26% and 67% for glass and brick, respectively), suggesting that substratum type changes the efficiency of algal removal by brushing. Individual substrata and microsubstrata characteristics can have a strong effect on benthic algae development and potentially affect reach scale algal variability as mediated by geomorphology.     

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.     

Development of filamentous green algae in the benthic algal community in a littoral sand-beach zone of Lake Biwa

Temporal changes of biomass and dominant species in benthic algal communities were investigated in a littoral sand-beach zone in the north basin of Lake Biwa from December 1999 to September 2000. Chlorophyll-a amounts of benthic algal communities per unit area of the sandy sediments rapidly increased from late April to June. Increases in biomass of the benthic algal communities are considered to result from the propagation of filamentous green algae Oedogonium sp. and Spirogyra sp. The cell numbers of filamentous green algae and chlorophyll-a amounts of benthic algal communities at depths of 30 and 50cm at a station protected by a breakwater in May were significantly higher than those of a station exposed directly to wave activity. Thus, the biomass accumulation of the benthic algal communities seems to be regulated strongly by wave disturbance. The development of filamentous green algae may contribute to the increase in biomass of the benthic algal community and to the changes in seasonal patterns of biomass in the sand-beach zone of Lake Biwa. We consider that the development of the filamentous green algal community in the littoral zone of Lake Biwa is the result of eutrophication.     

Field ecology of freshwater macroalgae in pools and ditches,with special attention to eutrophication

A review is presented on the occurrence of macroalgae in dune pools, ditches and moorland pools in The Netherlands, which were studied in the period 1980–1987. There are clear effects of eutrophication on biomass as well as on species composition. A general trend is that under mesotrophic or moderately eutrophic conditions, the filamentous macroalgae form a diversified component of the primary producers in shallow water bodies. Especially spore forming genera asSpirogyra andOedogonium can be represented with up to about 20 species per site. In hardwater ditches alsoVaucheria is an important component, especially the sediment dwellingVaucheria dichotoma. Under distinctly eutrophic conditions dense floating algal masses contain only one or a few species asCladophora glomerata, Enteromorpha intestinalis, Hydrodictyon reticulatum. Under hypertrophic conditions the water surface is covered by dense blanketing mats of duckweeds in summer and autumn. Besides trophic factors, other factors as alkalinity and chloride concentration play a role in species composition. The results are discussed in the framework of water quality assessment of polder waters and possible applications for water purification or use as manuring substance. The need for experimental ecophysiological studies is underlined.     

Comparing Effects of Nutrients on Algal Biomass in Streams in Two Regions with Different Disturbance Regimes and with Applications for Developing Nutrient Criteria

Responses of stream algal biomass to nutrient enrichment were studied in two regions where differences in hydrologic variability cause great differences in herbivory. Around northwestern Kentucky (KY) hydrologic variability constrains invertebrate biomass and their effects on algae, but hydrologic stability in Michigan (MI) streams permits accrual of high herbivore densities and herbivory of benthic algae. Multiple indicators of algal biomass and nutrient availability were measured in 104 streams with repeated sampling at each site over a 2−month period. Many measures of algal biomass and nutrient availability were positively correlated in both regions, however the amount of variation explained varied with measures of biomass and nutrient concentration and with region. Indicators of diatom biomass were higher in KY than MI, but were not related to nutrient concentrations in either region. Chl a and % area of substratum covered by Cladophora were positively correlated to nutrient concentrations in both regions. Cladophora responded significantly more to nutrients in MI than KY. Total phosphorus (TP) and total nitrogen (TN) explained similar amounts of variation in algal biomass, and not significantly more variation in biomass than dissolved nutrient concentrations. Low N:P ratios in the benthic algae indicated N as well as P may be limiting their accrual. Most observed responses in benthic algal biomass occurred in nutrient concentrations between 10 and 30 μg TP l⁻¹ and between 400 and 1000 μg TN l⁻¹.     

Recolonization of coastal heath by Pseudomys novaehollandiae (Muridae) following sand mining

The regeneration of coastal heath after disturbance by mineral sand mining was studied on mid-seral stages from 4 to 11 years old at Hawks Nest, N.S.W. The main purpose was to gain some understanding of factors influencing recolonization by pioneer species of small mammals such as Pseudomys novaehollandiae. Changes in floristics and vegetation structure with time were studied as possible contributing factors together with environmental variables. An apparently linear relationship was found between plant species diversity (X₁) and P. novaehollandiae biomass which was also correlated with an index representing the proportion of heath species present (X₂). Both plant species diversity and P. novaehollandiae biomass showed a linear increase with regeneration age. A multiple regression analysis revealed a predictive equation explaining 96% of the variation in P. novaehollandiae biomass (Y): Y=-7.92 + 1.21X₁+3.92X₂ - 3.09X₃ The third variable (X₃) is a measure of soil hardness. A path diagram using contribution coefficients based on a partial correlation analysis included the effects of vegetation structure below 50 cm and regeneration age. On mid-seral stages after sand mining P. novaehollandiae is associated with areas having a variety of heath plants, with vegetation cover below 50 cm and softer substrates; its abundance increases with increasing regeneration age and with the total amount of vegetation present. A minimum of 20 years is seen to be necessary for both total amount of vegetation and P. novaehollandiae biomass to reach values encountered on control plots of undisturbed heath.     

Reappraisal ofDiplolabellum coreanum (orchidaceae) as inferred from molecular data

The taxonomic treatment of a monotypic genus,Diplolabellum, has been disputed by various authors. Maekawa (1935) described the genus on the basis ofOreorchis coreana IFinet. Because distribution of its plants is very limited, i.e., to Jeju Island of the South Korean Peninsula, it has not been well studied. To reappraise the phylogenetic relationship ofDiplolabellum coreanum Finet, we obtained ITS,matK, trnT-trnL, andtrnL-trnF sequences from several species ofOreorchis and related genera. Sequence analysis showed thatD. coreanum is closely related to one group ofOreorchis that consists ofO. patens Lindle andO. fargesii Finet. Therefore, our molecular data support treating the species asO. coreana rather than asD. coreanum, even though the latter genus is distinct fromOreorchis in morphological characters such as callus, pedicel, column, and caudicle.     

Biological,chemical and physical factors influencing the seasonal change of polysaccharide concentration in irrigation-channel sediment

Variation of polysaccharide concentration in irrigation-channel sediment was determined concurrently with biological, chemical and physical factors influencing the benthic algal community. Phenol-sulphuric acid method was used to measure polysaccharide concentration. Polysaccharide concentration, biomass of benthic algae, and species composition changed spatially and temporally. Fluctuations of total suspended solid (TSS) concentration and exposure of channel bed to direct sunlight had major effects on algal growth and polysaccharide production. Polysaccharide concentration was correlated to chlorophyll a concentration (r=0.73, P<0.001) and algal biomass (r=0.57, P<0.001). Fragilaria construens and Aulacoseira (Melosira) italica were the most common diatoms in the benthic flora. Chlorophyll a concentration in the sediment showed a strong negative correlation (r=-0.99, P<0.001) with the seasonal variation of TSS concentration in channel water. The polysaccharides produced by benthic microorganisms play a major role in clogging channel bed and thereby reducing seepage from earthen irrigation channels. Correlations between polysaccharide concentration and chlorophyll a (and algal biomass) further indicate the importance of benthic algae for polysaccharide production. Since availability of light to the algal flora is critical for the production of polysaccharides, the effect of clogging can be maximized by exposing the channel bed to direct sunlight during non-irrigation period (winter).     

Effects of stream predator richness on the prey community and ecosystem attributes

It is important to understand the role that different predators can have to be able to predict how changes in the predator assemblage may affect the prey community and ecosystem attributes. We tested the effects of different stream predators on macroinvertebrates and ecosystem attributes, in terms of benthic algal biomass and accumulation of detritus, in artificial stream channels. Predator richness was manipulated from zero to three predators, using two fish and one crayfish species, while density was kept equal (n = 6) in all treatments with predators. Predators differed in their foraging strategies (benthic vs. drift feeding fish and omnivorous crayfish) but had overlapping food preferences. We found effects of both predator species richness and identity, but the direction of effects differed depending on the response variable. While there was no effect on macroinvertebrate biomass, diversity of predatory macroinvertebrates decreased with increasing predator species richness, which suggests complementarity between predators for this functional feeding group. Moreover, the accumulation of detritus was affected by both predator species richness and predator identity. Increasing predator species richness decreased detritus accumulation and presence of the benthic fish resulted in the lowest amounts of detritus. Predator identity (the benthic fish), but not predator species richness had a positive effect on benthic algal biomass. Furthermore, the results indicate indirect negative effects between the two ecosystem attributes, with a negative correlation between the amount of detritus and algal biomass. Hence, interactions between different predators directly affected stream community structure, while predator identity had the strongest impact on ecosystem attributes.     

Effects of eutrophication on the interaction between algae and grazers in an Andean stream

Nutrient excess is a common disturbance that affects biological interactions in river ecosystems. The response of nutrient supply on primary producers and Tricorythodes sp., a common mayfly grazer, was determined in experimental chambers set in a tropical, high Andean stream. Chambers in an experimentally fertilized reach developed higher amount of both benthic and detached chlorophyll than chambers in an upstream control reach. Fertilization produced a slight increase in grazer biomass, and reduced algal biomass compared to grazer-free chambers. These results show that nutrient excess spread bottom-up effects through the food web, and that relevant top-down effects could also be detected. Eutrophication may produce relevant changes in the food web of tropical high-mountain streams.     

Food habits of the farmer damselfish <Emphasis Type="Italic">Stegastes nigricans</Emphasis> inferred by stomach content,stable isotope,and fatty acid composition analyses

The territorial damselfish, Stegastes nigricans, maintains algal farms by excluding invading herbivores and weeding unpalatable algae from its territories. In Okinawa, Japan, S. nigricans farms are exclusively dominated by Polysiphonia sp., a highly digestible filamentous rhodophyte. This study was aimed at determining the diet of S. nigricans in Okinawa and its dependency on these almost-monoculture algal farms based on stomach content and chemical analyses. Stomach content analyses revealed that all available food items in the algal farms (i.e., algae, benthic animal inhabitants, trapped detritus) were contained in fish stomachs, but amorphous organic matter accounted for 68% of the contents. Therefore, carbon and nitrogen stable isotope ratios and fatty acid (FA) compositions were analyzed to trace items actually assimilated in their bodies. Stable isotope analyses showed that benthic animals were an important food source even for this farmer fish. Two essential fatty acids (EFAs), 20:4n6 and 20:5n3, which are produced only by rhodophytes among available food items, were rich in the muscle tissue of S. nigricans as well as in algal mats and detritus, suggesting that algal mats contribute EFAs to S. nigricans directly and indirectly through the food web. In conclusion, S. nigricans ingested algal mats, detritus, and benthic animals maintained within its farm. Algae and detritus were original sources of EFAs, and benthic animals, which were much more abundant in the farms than in outside territories, provided a nitrogen-rich dietary source for the fish.     

Gastropod grazers and nutrients,but not light,interact in determining periphytic algal diversity

The potential interactions of grazing, nutrients and light in influencing autotroph species diversity have not previously been considered. Earlier studies have shown that grazing and nutrients interact in determining autotroph species diversity, since grazing decreases species diversity when nutrients (i.e. N or P) limit autotroph growth, but increases it when nutrients are replete. We hypothesized that increased light intensities would intensify the interactions between grazing and nutrients on algal species diversity, resulting in even stronger reductions in algal species diversity through grazing under nutrient–poor conditions, and to even stronger increases of algal species diversity through grazing under nutrient-rich conditions. We studied the effects of grazing (absent, present), nutrients (ambient, N + P enriched) and light (low light, high light) on benthic algal diversity and periphyton C:nutrient ratios (which can indicate algal nutrient limitation) in a factorial laboratory experiment, using the gastropod grazer Viviparus viviparus. Grazing decreased algal biomass and algal diversity, but increased C:P and N:P ratios of periphyton. Grazing also affected periphyton species composition, by decreasing the proportion of Spirogyra sp. and increasing the proportion of species in the Chaetophorales. Grazing effects on diversity as well as on periphyton N:P ratios were weakened when nutrients were added (interaction between grazing and nutrients). Chlorophyll a (Chl a) per area increased with nutrient addition and decreased with high light intensities. Light did not increase the strength of the interaction between grazing and nutrients on periphytic algal diversity. This study shows that nutrient addition substantially reduced the negative effects of grazing on periphytic algal diversity, whereas light did not interact with grazing or nutrient enrichment in determining periphytic algal diversity.     

Current velocity influences the facilitation and removal of algae by stream grazers

The modification of flows in lotic ecosystems can have dramatic effects on abiotic and biotic processes and change the structure of basal trophic levels. In high-gradient streams, most of the biota are benthic, and decreased flow may homogenize and reduce benthic current velocity, potentially changing stream ecosystem function. Grazing by macroinvertebrates is an important component of stream function because grazers regulate energy flow from primary producers to higher trophic levels. We conducted an experiment to examine how macroinvertebrate grazers facilitated or removed algal biomass across a gradient of benthic current velocity (0–40 cm s^?1). We chose three grazers (Drunella coloradensis, Cinygmula spp., and Epeorus deceptivus) from a montane stream and conducted our experiment using 24 artificial stream channels that had three treatments: no grazers (control), single-grazer, and combined-grazer treatments. In the absence of grazers, algal biomass increased with benthic current velocity. Grazer treatments differed from the control in that more algal biomass was removed at higher velocities, whereas algal accrual was largely facilitated at low velocities. The transition from facilitation to removal ranged from 4.5 to 5.9 cm s^?1 for individual grazer treatments and occurred at 11.7 cm s^?1 for the combined-grazer treatment. Our data suggest that velocity plays a significant role in the facilitation and removal of algae by macroinvertebrate grazers. Additionally, the patterns revealed here could have general implications for algal accrual in systems where flow is reduced.     

The effect of two scales of habitat architecture on benthic grazing in a river

1. This experiment studied the effects of differing levels of the complexity of substratum architecture at two spatial scales on the distribution and abundance of benthic algae and invertebrates, and the strength of the trophic interaction between invertebrate grazers and algae. Some estimates of the effects on invertebrate colonization rates were also made. 2. Four levels of microhabitat architectural complexity were created using artificial substrata (clay tiles) and placed in Mountain River, Tasmania, in two riffle types (bedrock and boulder-cobble) of differing large-scale substratum complexity. After a colonization period, invertebrate grazers were removed from half the tiles to measure the effects of grazing. Invertebrates on the tiles were also counted and identified. At the end of the experiment, algae were removed from the tiles and analysed for chlorophyll a. 3. Invertebrate grazers did not reduce algal biomass during the experiment, and microhabitat-scale architecture influenced algal biomass more strongly than riffle-scale architecture. Highly complex microhabitat architecture increased algal biomass by providing more surface area, but once standardized for surface area, algal biomass decreased as the complexity of microhabitat architecture increased. 4. Microhabitat-scale architecture was also predominant in determining invertebrate density and the identity of the dominant grazer species. In contrast to algal biomass, invertebrate densities and species density increased with the complexity of microhabitat architecture, suggesting that refuges from flow (and possibly predation) were as important to river invertebrates as the distribution of their food source. 5. Riffle-scale architecture had some effect on the colonization of two slow-moving grazer taxa, but, overall, the colonization processes of slow-moving grazers were determined mostly by the complexity of microhabitat-scale architecture.     

Herbivores control effects of algal species richness on community biomass and stability in a laboratory microcosm experiment

Hundreds of studies that have explored how biodiversity affects the productivity and stability of ecosystems have produced a consensus that communities composed of more species tend to have higher biomass that is more stable through time. However, the majority of this work stems from studies performed using highly simplified food webs, often composed of just primary producers competing for inorganic resources in the absence of trophic interactions. When studies have incorporated trophic interactions, diversity‐function relationships have been more variable, leaving open the question of how biodiversity affects the functioning of ecosystems with more trophic levels. Here we report the results of a laboratory experiment that used freshwater microcosms to test for effects of algal diversity (one or four species) on community biomass and temporal variability in the presence and absence of two different herbivore species (cladocerans Ceriodaphnia dubia and Daphnia pulex). When no herbivores were present, we found the classic pattern observed in hundreds of other studies – as species richness of algae increased, algal biomass increased, and the temporal variation in biomass decreased. This pattern was retained when one of the herbivores (C. dubia) was present. Ceriodaphnia dubia exhibited weak and non‐selective grazing on the focal algae, leaving the effect of diversity on biomass and variability essentially intact. In contrast, D. pulex exhibited strong and selective grazing in algal polycultures that qualitatively altered both diversity–function relationships. As algal richness increased, total algal biomass decreased and variation through time increased. These changes were coupled with larger and less variable populations of D. pulex. Our results show that herbivory leads to a richer array of diversity–function relationships than often observed in studies focused on just one trophic level, and suggests trophic interactions should be given more attention in work that seeks to determine how biodiversity impacts the functioning of ecosystems.     

Catchment urbanisation and increased benthic algal biomass in streams: linking mechanisms to management

1. Urbanisation is an important cause of eutrophication in waters draining urban areas. We determined whether benthic algal biomass in small streams draining urban areas was explained primarily by small‐scale factors (benthic light, substratum type and nutrient concentrations) within a stream, or by catchment‐scale variables that incorporate the interacting multiple impacts of urbanisation (i.e. variables that describe urban density and the intensity of drainage or septic tank systems). 2. Benthic algal biomass was assessed as chlorophyll a density (chl a) in 16 streams spanning a rural–urban gradient, with both a wide range of urban density and of piped stormwater infrastructure intensity on the eastern fringe of metropolitan Melbourne, Australia. The gradient of urban density among streams was broadly correlated with catchment imperviousness, drainage connection (proportion of impervious areas connected to streams by stormwater pipes), altitude, longitude and median phosphorus concentration. Catchment area, septic tank density, median nitrogen concentration, benthic light (photosynthetically active radiation) and substratum type were not strongly correlated with the urban gradient. 3. Variation in benthic light and substratum type within streams explained a relatively small amount of variation in log chl a (3–11 and 1–13%, respectively) compared with between‐site variation (39–54%). 4. Median chl a was positively correlated with catchment urbanisation, with a large proportion of variance explained jointly (as determined by hierarchical partitioning) by those variables correlated with urban density. Independent of this correlation, the contributions of drainage connection and altitude to the explained variance in chl a were significant. 5. The direct connection of impervious surfaces to streams by stormwater pipes is hypothesised as the main determinant of algal biomass in these streams through its effect on the supply of phosphorus, possibly in interaction with stormwater‐related impacts on grazing fauna. Management of benthic algal biomass in streams of urbanised catchments is likely to be most effective through the application of stormwater management approaches that reduce drainage connection.     

Sources of organic carbon in the littoral of Lake Gooimeer as indicated by stable carbon isotope and carbohydrate compositions

The relative importance of potential carbon sources in the littoral of Lake Gooimeer, a lake in the centre of the Netherlands, was studied using a combination of ¹³C/¹²C-ratio analysis and carbohydrate composition analysis. The littoral is covered on the land side by a 80 m wide Phragmites australis bed. Potential carbon sources were macrophyte litter, seston and benthic algae. Samples of potential carbon sources, sediments and benthic macrofauna from inside and outside the bed were analyzed for their¹³C/¹²C-ratio and some for their carbohydrate composition. Results indicate that inside the bed, macrophyte litter was the main source of carbon for both the sediment organic matter and the benthic macrofauna, and that algal material was of minor importance. Outside the bed, production by benthic algae was the main carbon source, with seston as a second source. No macrophyte derived material could be detected outside the reed bed.Abbreviations DOC (Dissolved Organic Carbon) - SOM (Sediment Organic Matter)