Laurentian Great Lakes Phytoplankton and Their Water Quality Characteristics,Including a Diatom-Based Model for Paleoreconstruction of Phosphorus

Recent shifts in water quality and food web characteristics driven by anthropogenic impacts on the Laurentian Great Lakes warranted an examination of pelagic primary producers as tracers of environmental change. The distributions of the 263 common phytoplankton taxa were related to water quality variables to determine taxon-specific responses that may be useful in indicator models. A detailed checklist of taxa and their environmental optima are provided. Multivariate analyses indicated a strong relationship between total phosphorus (TP) and patterns in the diatom assemblages across the Great Lakes. Of the 118 common diatom taxa, 90 (76%) had a directional response along the TP gradient. We further evaluated a diatom-based transfer function for TP based on the weighted-average abundance of taxa, assuming unimodal distributions along the TP gradient. The r² between observed and inferred TP in the training dataset was 0.79. Substantial spatial and environmental autocorrelation within the training set of samples justified the need for further model validation. A randomization procedure indicated that the actual transfer function consistently performed better than functions based on reshuffled environmental data. Further, TP was minimally confounded by other environmental variables, as indicated by the relatively large amount of unique variance in the diatoms explained by TP. We demonstrated the effectiveness of the transfer function by hindcasting TP concentrations using fossil diatom assemblages in a Lake Superior sediment core. Passive, multivariate analysis of the fossil samples against the training set indicated that phosphorus was a strong determinant of historical diatom assemblages, verifying that the transfer function was suited to reconstruct past TP in Lake Superior. Collectively, these results showed that phytoplankton coefficients for water quality can be robust indicators of Great Lakes pelagic condition. The diatom-based transfer function can be used in lake management when retrospective data are needed for tracking long-term degradation, remediation and trajectories.     

Water quality assessment using diatom assemblages and advanced modelling techniques

1. Two types of artificial neural networks procedures were used to define and predict diatom assemblage structures in Luxembourg streams using environmental data. 2. Self‐organising maps (SOM) were used to classify samples according to their diatom composition, and multilayer perceptron with a backpropagation learning algorithm (BPN) was used to predict these assemblages using environmental characteristics of each sample as input and spatial coordinates (X and Y) of the cell centres of the SOM map identified as diatom assemblages as output. Classical methods (correspondence analysis and clustering analysis) were then used to identify the relations between diatom assemblages and the SOM cell number. A canonical correspondence analysis was also used to define the relationship between these assemblages and the environmental conditions. 3. The diatom‐SOM training set resulted in 12 representative assemblages (12 clusters) having different species compositions. Comparison of observed and estimated sample positions on the SOM map were used to evaluate the performance of the BPN (correlation coefficients were 0.93 for X and 0.94 for Y). Mean square errors of 12 cells varied from 0.47 to 1.77 and the proportion of well predicted samples ranged from 37.5 to 92.9%. This study showed the high predictability of diatom assemblages using physical and chemical parameters for a small number of river types within a restricted geographical area.     

COMPARISON OF SIMPLE AND MULTIMETRIC DIATOM‐BASED INDICES FOR GREAT LAKES COASTLINE DISTURBANCE1

Because diatom communities are subject to the prevailing water quality in the Great Lakes coastal environment, diatom‐based indices can be used to support coastal‐monitoring programs and paleoecological studies. Diatom samples were collected from Great Lakes coastal wetlands, embayments, and high‐energy sites (155 sites), and assemblages were characterized to the species level. We defined 42 metrics on the basis of autecological and functional properties of species assemblages, including species diversity, motile species, planktonic species, proportion dominant taxon, taxonomic metrics (e.g., proportion Stephanodiscoid taxa), and diatom‐inferred (DI) water quality (e.g., DI chloride [Cl]). Redundant metrics were eliminated, and a diatom‐based multimetric index (MMDI) to infer coastline disturbance was developed. Anthropogenic stresses in adjacent coastal watersheds were characterized using geographic information system (GIS) data related to agricultural and urban land cover and atmospheric deposition. Fourteen independent diatom metrics had significant regressions with watershed stressor data; these metrics were selected for inclusion in the MMDI. The final MMDI was developed as the weighted sum of the selected metric scores with weights based on a metric’s ability to reflect anthropogenic stressors in the adjacent watersheds. Despite careful development of the multimetric approach, verification using a test set of sites indicated that the MMDI was not able to predict watershed stressors better than some of the component metrics. From this investigation, it was determined that simpler, more traditional diatom‐based metrics (e.g., DI Cl, proportion Cl‐tolerant species, and DI total phosphorus [TP]) provide superior prediction of overall stressor influence at coastal locales.     

FRESHWATER DIATOMS FROM THE CANADIAN ARCTIC TREELINE AND DEVELOPMENT OF PALEOLIMNOLOGICAL INFERENCE MODELS1

Relationships between surface sediment diatom assemblages and measured environmental variables from 77 lakes in the central Canadian arctic treeline region were examined using multivariate statistical methods. Lakes were distributed across the arctic treeline from boreal forest to arctic tundra ecozones, along steep climatic and environmental gradients. Forward selection in canonical correspondence analysis determined that dissolved inorganic carbon (DIC), dissolved organic carbon (DOC), total nitrogen (TN), lake surface area, silica, lake‐water depth, and iron explained significant portions of diatom species variation. Weighted‐averaging (WA) regression and calibration techniques were used to develop inference models for DIC, DOC, and TN from the estimated optima of the diatom taxa to these environmental variables. Simple WA models with classical deshrinking produced models with the strongest predictive abilities for all three variables based on the bootstrapped root mean squared errors of prediction (RMSEP). WA partial least squares showed little improvement over the simpler WA models as judged by the jackknifed RMSEP. These models suggest that it is possible to infer trends in DIC, DOC, and TN from fossil diatom assemblages from suitably chosen lakes in the central Canadian arctic treeline region.     

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.     

Regional distribution of diatom assemblages in the headwater streams of Luxembourg

The aim of this study was to determine the influence of environmental variables on the structure of benthic diatom assemblages, and to propose type assemblages of diatoms characterizing unpolluted headwater streams of Luxembourg. A total of 289 diatom samples were collected in the headwater streams of Luxembourg. At each sampling site, physical and chemical variables were also measured. The relationships between environmental variables and the distribution of the taxa were estimated using univariate and multivariate statistical analyses. On the basis of geology and stream water chemistry, Luxembourg could be separated in two regions. The streams in the schistose northern part of Luxembourg presented a low carbonate hardness (median 3.5° F); the streams in the southern part of Luxembourg presented a high carbonate hardness (median 21.8° F) notably connected to the presence of sandstone and limestone substrata. A Twinspan classification carried out on diatom assemblages defined two groups of samples corresponding to these two regions. The carbonate hardness that is related to the nature of the geological substratum appeared to be the major structuring variable for the assemblage composition. Anthropogenic pollution was a secondary structuring variable for diatom assemblages since each group could be subdivided in subgroups presenting statistically different nutrient and organic matter concentrations. The diatom assemblages of these subgroups were characterised by differences of saprobic and trophic preferences. On the basis of these results, two type assemblages of diatoms are proposed for the unpolluted headwater streams in the two regions of Luxembourg. The presence of these different type assemblages in Luxembourg shows the necessity to adapt diatom bioindication to the different regions of the country.     

Prediction of benthic community structure from environmental variables in a soft-sediment tidal basin (North Sea)

The relationship between benthos data and environmental data in 308 samples collected from the intertidal zone of the H?rnum tidal basin (German Wadden Sea) was analyzed. The environmental variables were current velocity, wave action, emersion time (all of which were obtained from a 2-year simulation with a numerical model) and four sediment grain-size parameters. A grouping of sample stations into five benthos clusters showed a large-scale (>1?km) zoning of benthic assemblages on the tidal flats. The zoning varied with the distance from the shore. Three sample applications were examined to test the predictability of the benthic community structure based on environmental variables. In each application, the dataset was spatially partitioned into a training set and a test set. Predictions of benthic community structure in the test sets were attempted using a multinomial logistic regression model. Applying hydrodynamic predictors, the model performed significantly better than it did when sediment predictors were applied. The accuracy of model predictions, given by Cohen’s kappa, varied between 0.14 and 0.49. The model results were consistent with independently attained evidence of the important role of physical factors in Wadden Sea tidal flat ecology.     

DIATOM ASSEMBLAGES AS INDICATORS OF LAKE TROPHIC STATUS IN SOUTHEASTERN ONTARIO LAKES1

Canonical correspondence analysis (CCA) was used to explore the relationship between measured environmental variables and surficial diatom (Bacillariophyceae) assemblages in alkaline lakes from southeastern Ontario. Total nitrogen (TN), watershed area, alkalinity, and maximum depth each explain significant (P ≤ 0.05) directions of variance in the distribution of diatom taxa. TN was highly correlated to total phosphorus (TP) (r = 0.92), chlorophyll a (r = 0.86), and Secchi depth (r =0.77). When a series of CCAs were run with the first axis constrained to each of these variables in turn, the ratio of the eigenvalue of the first axis to that of the second axis (λ1/λ2) was highest for TN, indicating that TN best explained the distribution of the diatom assemblages in this set of lakes. Furthermore, results of Monte Carlo permutation tests indicated that these four variables did not act independently on the diatom assemblages. Therefore, TN was selected to represent these four closely related variables to infer lake trophic status. Weighted-averaging regression and calibration (with classical deshrinking) were used to develop transfer functions to infer TN from the relative abundances of 83 diatom taxa recovered from the surficial sediments of 51 lakes. There was a good correlation between diatom-inferred TN concentrations and measured TN concentrations (r²= 0.75, n = 51). The weighted-averaging regression and calibration model was used to infer lake trophic status (represented by TN) from diatom assemblages presented in the sediments from Little Round Lake, Ontario. These data were used in conjunction with historical land-use data in order to quantify the sequence and extent of nutrient enrichment related to human activity in the watershed area.     

Shifts in taxonomical and guild composition of littoral diatom assemblages along environmental gradients

Diatoms are commonly and frequently used as water quality indicators, but only a few studies have been done to evaluate the importance of littoral, contemporary diatoms as bioindicators. This study aims to determine the main predictors of diatom community composition from 73 Swedish lakes. Canonical correspondence analysis (CCA) revealed pH, phosphate, nitrite/nitrate levels, longitude and percentage of forest in the catchment to be the most important factors of 51 environmental variables for structuring diatom assemblages. Cluster analysis separated the lakes into three groups based on the diatom community composition. Lakes belonging to these groups were characterised as: (1) acidic, nutrient-poor; (2) circumneutral, nutrient-poor and (3) alkaline, nutrient-rich, according to the results of a discriminant function analysis and dominant diatom taxa revealed by similarity percentage analysis. Ecological guilds according to growth morphology and the ability of nitrogen-fixation were assigned to all diatom taxa. All three lake groups exhibited a distinct guild composition. Nitrogen-fixing diatoms were found in nutrient-rich lakes, only. Our results indicate that taxonomical composition of littoral diatom assemblages can be applied in the assessment of nutrient and acidity status of Swedish lakes. Differences in distribution of the ecological guilds were connected to several environmental factors such as nutrients, light and grazing; their application in assessment of trophic status of lakes is therefore precarious.     

Nutrients exert a stronger control than climate on recent diatom communities in Esthwaite Water: evidence from monitoring and palaeolimnological records

1. A long‐term monitoring programme on phytoplankton and physicochemical characteristics of Esthwaite Water (England) that started in 1945 provides a rare opportunity to understand the effects of climate and nutrients on a lake ecosystem. 2. Monitoring records show that the lake experienced nutrient enrichment from the early 1970s, particularly after 1975, associated with inputs from a local sewage treatment plant, resulting in marked increases in concentration of soluble reactive phosphorus (SRP). Climatic variables, such as air temperature (AirT) and rainfall, exhibit high variability with increasing trends after 1975. 3. Diatom analyses of an integrated ²¹⁰Pb‐dated lake sediment core from Esthwaite Water, covering the period from 1945 to 2004, showed that fossil diatoms exhibited distinct compositional change in response to nutrient enrichment. 4. Redundancy analysis (RDA) based on diatom and environmental data sets over the past 60 years showed that the most important variables explaining diatom species composition were winter concentrations of SRP, followed by AirT, independently explaining 22% and 8% of the diatom variance, respectively. 5. Additive models showed that winter SRP was the most important factor controlling the diatom assemblages for the whole monitoring period. AirT had little effect on the diatom assemblages when nutrient levels were low prior to 1975. With the increase in nutrient availability during the eutrophication phase after 1975, climate became more important in regulating the diatom community, although SRP was still the major controlling factor. 6. The relative effects of climate and nutrients on diatom communities vary depending on the timescale. RDA and additive model revealed that climate contributed little to diatom dynamics at an annual or decadal scale. 7. The combination of monitoring and palaeolimnological records employed here offers the opportunity to explore how nutrients and climate have affected a lake ecosystem over a range of timescales. This dual approach can potentially be extended to much longer timescales (e.g. centuries), where long‐term, reliable observational records exist.     

A multi-proxy paleolimnological reconstruction of trophic state reference conditions for stratified carbonate-rich lakes in northern Germany

This study aims to identify reference conditions (nutrient status and diatom assemblages) as required by the European Water Framework Directive (WFD) for stratified, carbonate-rich lowland lakes with a large watershed area (watershed area to lake volume ratio (WV) > 1.5 km² 10⁻⁶ m⁻³) and a retention time (RT) from 0.1 to 10 years (Central Baltic Lake-Type 1, German Lake-Type 10) in European ecoregion 14. Diatoms, pollen and geochemistry were analysed from sediment cores of six lakes from northern Germany representing different subtypes of Lake-Type 10 (varying WV and RT) and covering the past 290–1,750 years. Historic total phosphorus levels were inferred using diatom-based transfer functions selected from a merged European data set and from optimised data sets identified with the moving-window approach. Pollen and geochemical proxies were used to identify occurrence and intensity of anthropogenic catchment usage. Lake trophic state reference conditions and associated diatom assemblages were identified for three of the six study lakes. In contrast, according to fossil pollen assemblages, two lakes were already strongly impacted by intensive catchment usage when the oldest investigated sediments were laid down. Thus, reference conditions of these already eutrophic lakes could not be identified. Similarly, the lowermost samples of a core from the sixth lake showed signs of impact, and it remains unclear whether the identified dystrophic conditions occurred naturally or if they were due to the drainage of wetlands in Medieval times. Lakes with a relatively small WV (1.5–5.0 km² 10⁻⁶ m⁻³) and RT > 1 year were naturally oligotrophic to low mesotrophic and a typical, representative diatom assemblage was identified. In contrast, typical reference conditions or diatom assemblages for lakes with higher WV (5–18.6 km² 10⁻⁶ m⁻³) and RT < 1 year could not be identified as chemical precipitation and upstream lakes (nutrient sinks or sources) additionally influenced natural nutrient levels. Therefore, the reference situation of both trophic state and diatom assemblages in a lake may be strongly influenced by other modifying, limnological processes in addition to WV and RT. Overall, this study helps to implement the WFD by identifying reference conditions and by discussing the level of differentiation of lake types required to set reference conditions. Guest editors: K. Buczkó, J. Korponai, J. Padisák & S. W. Starratt Palaeolimnological Proxies as Tools of Environmental Reconstruction in Fresh Water     

The Role of Substrate Type on Benthic Diatom Assemblages in the Daly and Roper Rivers of the Australian Wet/Dry Tropics

The selection of one or more river substrata for the collection of benthic diatoms is fundamental to any monitoring or research programme because it can potentially bias the diatom data set. In the wet/dry tropics of Australia, where the use of benthic diatoms for river health assessment is in its infancy, the comparability of diatom assemblages on river substrata has been assessed. Benthic diatoms were sampled from seven river sites, with a range of ionic chemistries (conductivities 27–6500 μS cm⁻¹) but low nutrient concentrations. At each site, triplicate samples were collected from 3 to 6 substrata. The diatom assemblages sampled were: epilithon (assemblages on rock), epiphytes on macroalgae and macrophytes, epidendron (assemblages on wood), epipsammon (assemblages on sand), epipelon (assemblages on mud) and bacterial slime. The variability between substrate assemblages, at each site, were assessed according to the following: (1) a multivariate analysis of diatom assemblages, (2) ANOVA tests of species richness, (3) ANOVA tests of the relative abundance of common species (defined by an abundance of at least 10% in any one sample), and (4) a comparison of the number of species unique to a substrate. A total of 198 taxa were identified, with some taxa common to temperate Australia. Common species were found on all substrata, with sometimes statistically significantly different relative abundances. Taxa unique to a substrate had low relative abundances (0.1–2%), were most often found on only one replicate, and are unlikely to be substrate specific because many are known to occur on other substrata. The assemblages on hard substrata, epilithon and epidendron, were found to be most similar. Diatom assemblages on macroalgal and macrophyte substrata, compared to other substrata, were highly variable. This is attributed to the loss of diatoms from grazing and sloughing, followed by recolonisation of newly exposed substrate. Other assemblages, notably epipsammon, were similar to epilithon and epidendron but sometimes differed in their relative abundance of common species. The principal finding of the study was the similarity of the epilithon and epidendron, which are considered to be indistinguishable. Rock and wood hard substrata can be substituted for one another during field surveys, thereby increasing the number of potential sample sites available for monitoring activities that standardise to a hard substrate.     

Assessing the condition of the Missouri, Ohio, and Upper Mississippi rivers (USA) using diatom-based indicators

Diatom-based indicators were developed to assess environmental conditions in the Missouri, Ohio, and Upper Mississippi rivers. Disturbance gradients, comprising the first two principal components derived from a suite of stressor variables, included a trophic gradient (Trophic) and a gradient reflecting agriculture and other development activities (Ag/Dev). Diatom-based indicators were developed by creating models using weighted average calibration and regression-based transfer functions to relate planktonic and periphytic diatom species assemblages to each disturbance gradient. The most predictive disturbance models combined phytoplankton and periphyton assemblages into a single bioindicator model (observed versus inferred: Trophic $ r_{\text{boot}}^{2} = 0. 5 6 $ ; Ag/Dev $ r_{\text{boot}}^{2} = 0. 7 0 $ ). The geographic applicability of bioindicators was assessed by limiting sample geographical range during model calibrations. Geographic scale was limited by creating bioindicators using samples from: (a) each river, and (b) combined Mississippi/Missouri samples excluding Ohio River sites which were chemically unique. Indicator performance decreased with geographically restrictive models, therefore river basin-wide models, developed across all three rivers, is recommended. The most effective diatom-based disturbance bioindicators for this great river ecosystem could be applied using phytoplankton, periphyton, or combined assemblages to infer both trophic and agriculture/development disturbances.     

Modern diatom assemblages in Central Mexico: the role of water chemistry and other environmental factors as indicated by TWINSPAN and DECORANA

SUMMARY. 1. Modern diatom samples, for which physical and chemical environmental conditions were recorded, were collected from forty-seven sites across Central Mexico
2. The results of the diatom counts were analysed using the clustering program TWINSPAN and the ordination program DECORANA.
3. Results from 'Laguna Zacapu and associated sites' showed that where water chemistry varies little between sample sites the effects of habitat are emphasized. This data set highlighted the problems of looking for modern analogues where anthropogenic disturbance is great.
4. TWINSPAN results from the 'Other Sites' data set suggested possible changes in diatom assemblages with chemical composition. The DECORANA analyses, however, again seemed to emphasize habitat.
5. In the context of this study, bottom sediment samples appeared to provide the most useful picture of 'average' conditions and to be the best source of analogues for the interpretation of fossil assemblages.     

Are the landscape‐level drivers of water column and surface sediment diatoms different?

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RELATIONSHIPS BETWEEN LITTORAL DIATOMS AND THEIR CHEMICAL ENVIRONMENT IN NORTHEASTERN GERMAN LAKES AND RIVERS1

We explored statistical relationships between the composition of littoral diatom assemblages and 21 chemical and physical environmental variables in 69 lakes and 15 river sites in the lowland of northeastern Germany. Canonical correspondence analysis with single treatment and with forward selection of environmental variables was used to detect 11 important ecological variables (dissolved inorganic carbon [DIC], Na ⁺ , total phosphorus [TP], dissolved organic carbon [DOC], total nitrogen [TN], pH, oxygen saturation, dissolved iron, SO₄^{2 ?} , NH₄ ⁺ , soluble reactive silicium) and maximum water depth or Ca^{2 +} or soluble reactive phosphorus that most independently explain major proportions of the total diatom variance among the habitats. Monte Carlo permutation tests showed that each contributed a significant additional proportion (P < 0.05) of the variance in species composition. Together, these 11 most important environmental variables explained 34% of the total variance in species composition among the sites and captured 73% of the explained variance from the full 21 parameters model. Weighted‐averaging regression and calibration of 304 indicator taxa with tolerance down‐weighting and classic deshrinking was used to develop transfer functions between littoral diatoms and DIC, pH, TP, TN, and Cl ^? . The DOC:TP ratio was introduced and a weighted‐averaging model was developed to infer allochthonous DOC effects in freshwater ecosystems. This diatom‐DOC/TP model was significant (P < 0.001) and explained 7.6% of the total diatom variance among the sites, surpassing the inferential power of the diatom‐TP‐transfer function (7.3% explained variance). The root‐mean‐square errors of prediction of the models were estimated by jack‐knifing and were comparable with published data sets from surface sediment diatom samples. The data set of littoral diatoms and environmental variables allows use of the diatom‐environmental transfer functions in biomonitoring and paleolimnological approaches across a broad array of natural water resources (such as floodplains, flushed lakes, estuaries, shallow lakes) in the central European lowland ecoregion.     

NUTRIENT LIMITATION OF BENTHIC ALGAE IN LAKE MICHIGAN: THE ROLE OF SILICA1

In the Laurentian Great Lakes, phytoplankton growth and biomass are secondarily limited by silica (Si), as a result of phosphorus (P) enrichment. Even modest levels of P enrichment can induce secondary Silimitation, which, in turn, promotes a shift from the native diatom phytoplankton flora to chlorophyte and cyanobacteria species. However, very little is known about the nutritional status of benthic populations and their response to nutrient enrichment. Two experiments were performed in the littoral zone of Lake Michigan where nutrients were delivered to in situ benthic algal (episammic and epilithic) assemblages using nutrient‐diffusing substrata. In order to test the hypothesis that benthic algae in Lake Michigan are Si limited, a 2 × 3 factorial experiment was used to deliver all combinations of Si, N, and P to resident assemblages growing on artificial substrata composed of natural (Si rich) versus calcium carbonate (Si poor) sand. A second experiment utilized a serial enrichment to evaluate the role of Si in mediating changes in taxonomic composition. These findings indicate that benthic algae in Lake Michigan exhibit signs of secondary Si limitation, and that their response to enrichment is similar to the phytoplankton. Moreover, natural sand substrata may provide a source of Si to resident benthic algae.     

Epilithic diatom assemblages and their relationship to environmental characteristics in an agricultural watershed (Guadiana River,SW Spain)

Large-scale patterns of benthic diatom assemblages were analyzed in an agricultural basin, the Guadiana River. The distribution patterns of epilithic diatom assemblages were analyzed at different spatial scales: the whole watershed, the upper calcareous subcatchment and the mid-lower siliceous subcatchment. At the whole watershed scale, two major ecological gradients were revealed. The first one summarized the diatom distribution throughout a nutrient concentration gradient, while the second gradient was related to the geological structure of the watershed. Variance partitioning allowed the effects of the different sets of environmental parameters related to every CCA gradient to be separated. Analyzing the subcatchment gradients with partial CCA allowed us to define specific key factors that affect diatom species composition. Although water chemistry consistently played the most important role in structuring diatom assemblages in the Guadiana, spatial factors such as altitude or geographic location also explained some variation in diatom distribution.     

Can filter‐feeding Asian carp invade the Laurentian Great Lakes? A bioenergetic modelling exercise

1. There is much concern that filter‐feeding Asian carp will invade the Laurentian Great Lakes and deplete crucial plankton resources. We developed bioenergetic models, using parameters from Asian carp and other fish species, to explore the possibility that planktonic food resources are insufficient to support the growth of silver carp (Hypophthalmichthys molitrix) and bighead carp (H. nobilis) in the Great Lakes. 2. The models estimated basic metabolic requirements of silver and bighead carp under various body sizes, swimming speeds and reproductive stages. These requirements were then related to planktonic food resources and environmental temperature to predict when and where silver and bighead carp may survive in the Great Lakes, and how far they may travel. 3. Parameter values for respiration functions were derived experimentally in a coordinated study of silver and bighead carp, while consumption parameters were obtained from the literature on silver carp. Other model parameters lacking for Asian carp, such as those for egestion and excretion, were obtained from the literature on other fish species. 4. We found that full‐sized bighead carp required 61.0 kJ d^?1 just to maintain their body mass at 20 °C, approximately equivalent to feeding in a region with 255 μg L^?1 macrozooplankton (dry) or 10.43 μg L^?1 chlorophyll a. Silver carp energy requirements were slightly higher. 5. When applied to various habitats in the Great Lakes, our results suggest that silver and bighead carp will be unable to colonise most open‐water regions because of limited plankton availability. However, in some circumstances, carp metabolism at lower temperatures may be low enough to permit positive growth even at very low rations. Positive growth is even more likely in productive embayments and wetlands, and the modelled swimming costs in some of these habitats suggest that carp could travel >1 km d^?1 without losing biomass. 6. The simulation (and firmly hypothetical) results from this modelling study suggest when and where Asian carp could become established in the Great Lakes. Given the potential consequences to Great Lakes ecosystems if these filter feeders do prove capable of establishing reproducing populations, efforts to keep Asian carp out of the Great Lakes must not be lessened. However, we do encourage the use of bioenergetic modelling in a holistic approach to assessing the risk of Asian carp invasion in the Great Lakes region.     

THE INFLUENCE OF SUBMERGED MACROPHYTES ON SEDIMENTARY DIATOM ASSEMBLAGES1

Submerged macrophytes are a central component of lake ecosystems; however, little is known regarding their long‐term response to environmental change. We have examined the potential of diatoms as indicators of past macrophyte biomass. We first sampled periphyton to determine whether habitat was a predictor of diatom assemblage. We then sampled 41 lakes in Quebec, Canada, to evaluate whether whole‐lake submerged macrophyte biomass (BiomEpiV) influenced surface sediment diatom assemblages. A multivariate regression tree (MRT) was used to construct a semiquantitative model to reconstruct past macrophyte biomass. We determined that periphytic diatom assemblages on macrophytes were significantly different from those on wood and rocks (ANOSIM R = 0.63, P < 0.01). A redundancy analysis (RDA) of the 41‐lake data set identified BiomEpiV as a significant (P < 0.05) variable in structuring sedimentary diatom assemblages. The MRT analysis classified the lakes into three groups. These groups were (A) high‐macrophyte, nutrient‐limited lakes (BiomEpiV ≥525 μg · L^?1; total phosphorus [TP] <35 μg · L^?1; 23 lakes); (B) low‐macrophyte, nutrient‐limited lakes (BiomEpiV <525 μg · L^?1; TP <35 μg · L^?1; 12 lakes); and (C) eutrophic lakes (TP ≥35 μg · L^?1; six lakes). A semiquantitative model correctly predicted the MRT group of the lake 71% of the time (P < 0.001). These results suggest that submerged macrophytes have a significant influence on diatom community structure and that sedimentary diatom assemblages can be used to infer past macrophyte abundance.