Aim To study large-scale patterns of benthic diatom assemblages in rivers, to assess the relative importance of environmental and geographical factors affecting their composition, and to evaluate the implications of these patterns for the use of diatoms as indicators of water quality, particularly nutrient enrichment.Location The United States Geological Survey (USGS) National Water Quality Assessment (NAWQA) Programme data set covers the conterminous United States.
MethodsWe employed gradient analysis to understand the floristic structure of the data set and to discover major ecological gradients underlying variation in species composition at different spatial scales (entire US, Omernik Level 1 and Level 2 ecoregions). We used variance partitioning to separate the effect of environmental and spatial characteristics.Results At the national scale, three major ecological gradients were evident. The first was a complex 'downstream' gradient from fast-flowing, mostly oligotrophic, highland rivers to predominantly eutrophic rivers of high-and low-elevation plains. The second was a gradient integrating water mineral content and pH, which separated the soft and often more acidic waters of the humid eastern part of US from the alkaline waters of arid western regions. The third obvious gradient was related to latitudinal and altitudinal variation of temperature. Up to one-third of the total explainable variation in species data was attributed solely to geographical factors not correlated with measured environmental characteristics. We present several examples of species with complex patterns of spatial distribution.Main conclusions Although environment consistently plays the most important role in structuring diatom assemblages in rivers, spatial factors also explain some variation in diatom distribution, especially at the continental scale. Most of the species that are confined to limited geographical areas are not yet described and await future taxonomic work. We show that selection of species which could serve as indicators of nutrient enrichment in US rivers is not a straightforward procedure. The existence of complex environmental gradients, and still poorly understood spatial patterns of species distribution, precludes attempts to develop uniform diatom-based metrics that would be applicable everywhere in the US. We advocate the development and calibration of metrics based on data sets collected from more limited geographical areas, and that include sites having relatively narrow ranges of environmental characteristics other than that which the metrics are designed to indicate.
Summary
We quantified the relationships between diatom relative abundance and water conductivity and ionic composition, using a dataset of 3239 benthic diatom samples collected from 1109 river sites throughout the U.S.A. [U.S. Geological Survey National Water‐Quality Assessment (NAWQA) Program dataset]. This dataset provided a unique opportunity to explore the autecology of freshwater diatoms over a broad range of environmental conditions.
Conductivity ranged from 10 to 14 500 μS cm−1, but most of the rivers had moderate conductivity (interquartile range 180–618 μS cm−1). Calcium and bicarbonate were the dominant ions. Ionic composition, however, varied greatly because of the influence of natural and anthropogenic factors.
Canonical correspondence analysis (CCA) and Monte Carlo permutation tests showed that conductivity and abundances of major ions (HCO + CO, Cl−, SO, Ca2+, Mg2+, Na+, K+) all explained a statistically significant amount of the variation in assemblage composition of benthic diatoms. Concentrations of HCO + CO and Ca2+ were the most significant sources of environmental variance.
The CCA showed that the gradient of ionic composition explaining most variation in diatom assemblage structure ranged from waters dominated by Ca2+ and HCO + CO to waters with higher proportions of Na+, K+, and Cl−. The CCA also revealed that the distributions of some diatoms correlated strongly with proportions of individual cations and anions, and with the ratio of monovalent to divalent cations.
We present species indicator values (optima) for conductivity, major ions and proportions of those ions. We also identify diatom taxa characteristic of specific major‐ion chemistries. These species optima may be useful in future interpretations of diatom ecology and as indicator values in water‐quality assessment.
Variation of frustular morphology within the Achnanthidium minutissimum (Kütz.) Czarn. species complex was studied in type populations of 12 described taxa and in 30 recent North American river samples. The SEM observations in this study and other publications showed that ultrastructural characters on their own do not discriminate among taxa within the A. minutissimum complex. Therefore, an attempt was made to use other characters, such as valve shape and striation pattern, to delineate morphological groups. The sliding‐landmarks method was used to obtain valve‐shape descriptors. These shape variables were combined with conventional morphological characters in multivariate analyses. It was shown that some historically recognized taxa are morphologically distinct, while others are difficult to differentiate. Morphological grouping of “old” taxa most similar to A. minutissimum did not correspond to their taxonomic hierarchy in contemporary diatom floras. Morphometric analysis of a data set of 728 specimens from North American rivers revealed six morphological groups, although it was impossible to draw clear boundaries among them. These morphological groups differed significantly in their ecological characteristics and could be recommended as indicators of water quality. Application of the discriminant function analysis based on shape variables and striation pattern showed that North American specimens could be more consistently classified into the six groups identified in our analysis than into historically recognized taxa.
The large variation in size and shape in diatoms is shown by morphometric measurements of 515 benthic and pelagic diatom species from the Baltic Sea area. The largest mean cell dimension (mostly the apical axis) varied between 4.2 and 653 μm, cell surface area between 55 and 344,000 μm2, and cell volume between 21 and 14.2 × 106μm3. The shape‐related index, length to width ratio, was between 1.0 and 63.3 and the shape‐ and size‐related index, surface area to volume ratio, was between 0.02 and 3.13. Diatom community analysis by multivariate statistics is usually based on counts of a fixed number of diatom valves with species scores irrespective of cell size. This procedure underestimates the large species for two reasons. First, the importance of a species with higher cell volume is usually larger in a community. Second, larger species usually have lower abundances and their occurrence in the diatom counts is stochastic. This article shows that co‐occurring small and large diatom species can respond very differently to environmental constraints. Large epiphytic diatoms responded most to macroalgal host species and small epiphytic diatoms most to environmental conditions at the sampling site. Large epilithic diatoms responded strongly to salinity, whereas small epilithic diatoms did so less clearly. The conclusion is that different scale‐dependent responses are possible within one data set. The results from the test data also show that important ecological information from diatom data can be missed when the large species are neglected or underestimated.
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