The seasonal response of diatom communities to variable water quality in some English urban lakes

SUMMARY. 1. Five diatom assemblages and their corresponding site groups were associated with the urban lakes. When treated as a single dataset, DC A and PCA grouped these diatom assemblages along an environmental gradient from circum natural. dilute waters to strongly alkaline, concentrated, nutrient-rich waters.
2. When treated as individual datasets, 75% of the known variance was explained by the first axis of DCA in all five diatom assemblages; this indicated the existence of a strong primary environmental gradient. The PCA results showed that the identity of this primary environmental gradient can be different for each assemblage.
3. The statistical analyses showed that the physicochemical gradients were continuous and not discrete. Nevertheless, fine-scale analysis can identify smaller, more precise species and site groups.
4. Two types of species response could be identified within most sub-assemblages in the urban lakes: the first was a quasi-seasonal response and was characterized by a gradual rise to a peak relative abundance of one dominant or several co-dominant species which could be sustained for a period of weeks before a gradual decline. A quasi-seasonal response was sometimes interrupted by a storm response. The storm response was characterized by a sudden increase in the relative abundance of one or more diatom species following a storm, after which the species declined to pre-storm abundances within 7 days.
5. The effects of urban runoff on the water quality of urban lakes can override the effects of true seasonal factors such as water temperature and day length on the relative abundance of species.     

Freshwater diatom biogeography in the Canadian Arctic Archipelago

Aim Document the biogeographical distributions of diatoms in the Canadian Arctic in relation to environmental factors. Location The Canadian Arctic Archipelago. Methods Diatoms were extracted from lake sediments and treated using standard methods. Rarefaction‐estimated species richness, diatom concentrations (valves cc^?1), and diatom abundance were mapped using a Geographic Information System. The physical and chemical parameters of the lakes were measured. Results A total of 326 taxa from 63 genera were found in 62 lakes of the study area. Up to 85 and as low as eight taxa were identified in any one lake, and rarefaction‐estimated species richness correlated with lake size. Nearby lakes could have greatly different diatom assemblages. Many taxa showed limited distributions. Response surfaces and ordination techniques indicate that a large number of taxa prefer colder regions of the Arctic while the abundance of others was influenced by lake water chemical parameters such as TKN and SiO₂. Main conclusions Although many taxa were common and found across the study area, diatom assemblages showed regional differences in the Arctic. Some taxa were not found in either the northern or southern parts of the Archipelago and others were restricted to particular regions. Newly delineated genera showed interpretable geographical patterns and could be related to environmental factors, suggesting that this more natural grouping may enhance our understanding of diatom ecology. Geographical, physical, and chemical factors are needed to explain diatom distributions in the Arctic.     

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.     

PERIPHYTIC DIATOM ASSEMBLAGES FROM BATHURST ISLAND, NUNAVUT, CANADIAN HIGH ARCTIC: AN EXAMINATION OF COMMUNITY RELATIONSHIPS AND HABITAT PREFERENCES

Diatoms are potentially the most important biomonitors of environmental change in high arctic lakes and ponds, but to date few autecological data are available. Because of the shallow nature of many of these water bodies, a large proportion of taxa are periphytic and planktonic diatoms are absent for the most part. By determining the microhabitat and substrate preferences of these benthic diatom taxa, the potential exists to infer past changes in available habitats from fossil diatom assemblages collected from sediment cores and ultimately to reconstruct past environmental and climatic changes responsible for these shifts in habitat availability. To refine our understanding of high arctic diatom habitat preference, the common diatom taxa found on submerged moss (bryophyte), sediment, and rock substrates from lakes and ponds on Bathurst Island, Nunavut, Canadian High Arctic were examined. The relationships among key limnological variables and the common taxa from each habitat were examined. Many diatom taxa exhibited varying degrees of microhabitat preference, with moss representing the more unique habitat. In addition, the following limnological variables significantly ( P ≤ 0.05) explained the species variance for each of the three substrates: Na ⁺ and total nitrogen for moss; total phosphorus (filtered) and pH for rock; and Fe^{3 +} , total phosphorus (unfiltered), total nitrogen, temperature, and pH for sediment. These data can be used to help interpret monitoring and paleolimnological studies in this environmentally sensitive region.     

Climate drivers of diatom distribution in shallow subarctic lakes

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Paleolimnological investigation of three manipulated lakes from Sudbury,Canada

Stratigraphic changes in diatoms and chrysophytes from three manipulated Sudbury lakes were explored in an attempt to examine the influence of fertilization and/or neutralization on algal microfossil assemblages. Both diatom- and chrysophyte-inferred pH profiles indicate that the pH of Labelle Lake has remained fairly stable in the past. The study of Labelle and Middle lakes indicates that the addition of nutrients to acidic and non-acidic oligotrophic lakes did not directly influence diatom and chrysophyte species composition, perhaps because pH remained stable. The diatom and chrysophyte assemblages of Middle Lake only changed when the pH was raised. In Mountaintop Lake the recent shift in chrysophyte species composition and the resulting inferred pH decline is most likely related to a decline in mid-summer epilimnetic pH. Reliable paleolimnological inferences are difficult in lakes such as these because it is difficult to track limnological conditions in the absence of modern analogues.     

The ecology of the planktonic diatom Cyclotella and its implications for global environmental change studies

The fossil record of diatoms in lake sediments can be used to assess the effects of climate variability on lake ecosystems if ecological relationships between diatom community structure and environmental parameters are well understood. Cyclotella sensu lato taxa are a key group of diatoms that are frequently dominant members of phytoplankton communities in low‐ to moderate‐productivity lakes. Their relative abundances have fluctuated significantly in palaeolimnological records spanning over a century in arctic, alpine, boreal and temperate lakes. This suggests that these species are sensitive to environmental change and may serve as early indicators of ecosystem effects of global change. Yet patterns of change in Cyclotella species are not synchronous or unidirectional across, or even within, regions, raising the question of how to interpret these widespread changes in diatom community structure. We suggest that the path forward in resolving seemingly disparate records is to identify clearly the autecology of Cyclotella species, notably the role of nutrients, dissolved organic carbon and light, coupled with better consideration of both the mechanisms controlling lake thermal stratification processes and the resulting effects of changing lake thermal regimes on light and nutrients. Here we begin by reviewing the literature on the resource requirements of common Cyclotella taxa, illustrating that many studies reveal the importance of light, nitrogen, phosphorus, and interactions among these resources in controlling relative abundances. We then discuss how these resource requirements can be linked to shifts in limnological processes driven by environmental change, including climate‐driven change in lakewater temperature, thermal stratification and nutrient loading, as well as acidification‐driven shifts in nutrients and water clarity. We examine three case studies, each involving two lakes from the same region that have disparate trends in the relative abundances of the same species, and illustrate how the mechanisms by which these species abundances are changing can be deciphered. Ultimately, changes in resource availability and water clarity are key factors leading to shifts in Cyclotella abundances. Tighter integration of the autecology of this important group of diatoms with environmental change and subsequent alterations in limnological processes will improve interpretations of palaeolimnological records, and clarify the drivers of seemingly disparate patterns in fossil records showing widespread and rapid changes across the northern hemisphere.     

Diatom assemblages and their relationships to lakewater nitrogen levels and other limnological variables from 36 lakes and ponds on Banks Island,N.W.T., Canadian Arctic

Banks Island, in the Canadian Arctic Archipelago, has been identified as an important reference site for studies of environmental change, especially as it relates to climatic warming. The island is logistically manageable (i.e. researchers can survey the entire island in one field season) and, most importantly, spans three major ecoclimatic regions supporting a diverse and large bird and mammal population. Developing upon earlier work by the authors describing the limnology of Banks Island, this current study: (1) examines which physical and chemical limnological variables influence diatom assemblages in this relatively lush island; and (2) explores variations in the diatom assemblages by ecoclimatic zones. The relationship between diatom taxa from a 36 lake/pond surface sediment calibration set and a suite of limnological variables was explored using multivariate statistics. Dominant diatom species varied based on changing limnological characteristics, particularly between the colder, ultra-oligotrophic lakes in the more northern High Arctic regions compared to the warmer, more nitrogen-rich sites in the Low Arctic regions of Banks Island. Exploration of diatom ecoclimatic and environmental preferences revealed interesting relationships, including the development of a diatom-based transfer function that could be used to track overall trends on lakewater nitrogen concentrations, which may enable future paleolimnological studies to track shifts in nutrient levels and climatic, and other environmental changes. Handling editor: J. Padisak     

Lake regionalization and diatom metacommunity structuring in tropical South America

Lakes and their topological distribution across Earth's surface impose ecological and evolutionary constraints on aquatic metacommunities. In this study, we group similar lake ecosystems as metacommunity units influencing diatom community structure. We assembled a database of 195 lakes from the tropical Andes and adjacent lowlands (8°N–30°S and 58–79°W) with associated environmental predictors to examine diatom metacommunity patterns at two different levels: taxon and functional (deconstructed species matrix by ecological guilds). We also derived spatial variables that inherently assessed the relative role of dispersal. Using complementary multivariate statistical techniques (principal component analysis, cluster analysis, nonmetric multidimensional scaling, Procrustes, variance partitioning), we examined diatom–environment relationships among different lake habitats (sediment surface, periphyton, and plankton) and partitioned community variation to evaluate the influence of niche‐ and dispersal‐based assembly processes in diatom metacommunity structure across lake clusters. The results showed a significant association between geographic clusters of lakes based on gradients of climate and landscape configuration and diatom assemblages. Six lake clusters distributed along a latitudinal gradient were identified as functional metacommunity units for diatom communities. Variance partitioning revealed that dispersal mechanisms were a major contributor to diatom metacommunity structure, but in a highly context‐dependent fashion across lake clusters. In the Andean Altiplano and adjacent lowlands of Bolivia, diatom metacommunities are niche assembled but constrained by either dispersal limitation or mass effects, resulting from area, environmental heterogeneity, and ecological guild relationships. Topographic heterogeneity played an important role in structuring planktic diatom metacommunities. We emphasize the value of a guild‐based metacommunity model linked to dispersal for elucidating mechanisms underlying latitudinal gradients in distribution. Our findings reveal the importance of shifts in ecological drivers across climatic and physiographically distinct lake clusters, providing a basis for comparison of broad‐scale community gradients in lake‐rich regions elsewhere. This may help guide future research to explore evolutionary constraints on the rich Neotropical benthic diatom species pool.     

Correlation of surface sediment diatoms with the present lake water pH in 28 Algoma lakes,Ontario, Canada

The surface sediment diatom analysis of 28 Algoma lakes (pH 4.40–8.13) indicates that even though each lake has a widely different aquatic environment and characteristic diatom assemblage, a definite relationship exists between the lake water pH and their diatom assemblages. In the acidic lakes acidobiontic and acidophilous diatom species predominate whereas in circumneutral and alkaline lakes circumneutral and alkaliphilous diatoms were most common. Cluster analysis of the pH indicator diatom assemblages grouped the study lakes into three distinct cluster groups. These groups also closely corresponded to lake water pH. On the basis of published ecological information as well as their presence in our study lakes, the pH indicator status of a number of diatom taxa have been discussed. A detailed listing of the diatom taxa identified and their pH indicator status is provided in order to facilitate their use in future diatom-inferred pH studies.     

Sensitivity and responses of diatoms to climate warming in lakes heavily influenced by humans

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PERIPHYTIC DIATOM ASSEMBLAGES FROM ULTRA‐OLIGOTROPHIC AND UV TRANSPARENT LAKES AND PONDS ON VICTORIA ISLAND AND COMPARISONS WITH OTHER DIATOM SURVEYS IN THE CANADIAN ARCTIC1

Periphytic diatoms are potentially powerful indicators of environmental change in climatically‐sensitive high latitude regions. However, only a few studies have examined their taxonomic and ecological characteristics. We identified and enumerated diatom assemblages from sediment, rock, and moss habitats in 34 ultra‐oligotrophic and highly transparent lakes and ponds on Victoria Island, Arctic Canada. The similar limnological characteristics of the sites allowed us to examine the influence of habitat, independent of water chemistry, on the diatom assemblages. As is typical in shallow arctic water bodies, benthic taxa, including species of Achnanthes, Caloneis, Cymbella, Navicula, and Nitzschia, were most widely represented. Minor gradients in our measured environmental variables did not significantly explain any variance in diatom species, but there were marked differences in diatom assemblages among sites. Pond ephemerality seems to explain some diatom variation, because aerophilic taxa such as Achnanthes kryophila Petersen and A. marginulata Grunow were dominant in shallow sites that had undergone appreciable reductions in volume. We identified several taxa that exhibited strong habitat preferences to sediment, moss, or rock substrates and also found significant differences (P < 0.01) in diatom composition among the three habitats. In comparisons with three similar diatom surveys extending over 1200 km of latitude, we determined that surface sediment assemblages differed significantly (P < 0.001) among all regions examined. Diatom species diversity was inversely related to latitude, a result likely explained by differences in the lengths of growing seasons. These data contribute important ecological information on diatom assemblages in arctic regions and will aid in the interpretation of environmental changes in biomonitoring and paleolimnological studies.     

Diatom community response along a productivity gradient of shallow Antarctic lakes

Summary Antarctic lakes are simple ecosystems ranging in productivity from extremely low productive melt-water lakes to highly productive guano lakes. With the aim to assess the importance of environmental variables for species composition and abundance of periphytic diatoms, we quantified light availability and nutrient concentration in the water and in the sediment of 21 Antarctic lakes. Of a total of 66 diatom taxa identified, the distribution of 16 species were investigated in detail. Four of these (25%) showed highest abundance in highly productive lakes (low light/high nutrient availability), whereas three species (19%) increased in abundance as productivity decreased (high light/low nutrient availability). Some ecological theories predict that species richness shows a maximum in medium productive systems and a decreasing trend as productivity increases further due to competition. Other theories suggest that physical factors, including climate, is the major factor determining species richness. The results from our study suggest that none of the theories are completely wrong or right, but that they have to be combined to explain as much as possible of the variation in species richness.     

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.     

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.     

The palaeolimnological potential of diatom assemblages in floodplain lakes of the Danube Delta, Romania: a pilot study

Within the framework of a palaeolimnological pilot study, the potential of sedimentary diatom assemblages for the reconstruction of the eutrophication history was studied in short sediment cores from five shallow lakes located in the Romanian Danube Delta. A total of 234 diatom taxa representing 57 genera could be identified but diatom assemblages are generally predominated by a few species only. Loss on ignition percentages, total diatom abundances and diatom downcore successions largely vary within the single sediment cores and between all five lakes. This suggests the presence of various lake environments within the Danube Delta, each characterized by lake-specific geomorphological, sedimentological, hydrological, and biological conditions. Highly variable depositional conditions are also confirmed by radiocesium measurements in two of the five cores suggesting distinct inter-lake differences in the sedimentation rate. Causes for the great intra-lake and inter-lake variability in diatom downcore successions are difficult to specify and may include changes in the physico-chemical conditions, aquatic vegetation cover, water depth, river influence, turbidity and inter-biotic interactions. However, based on this pilot study, it is likely that the spatial and temporal distribution of sedimentary diatom assemblages in Danube Delta lakes were not triggered by the trophic state. The delta lakes likely became meso- to eutrophic long before 1950, possibly during late-Holocene times.     

Terrestrial and aquatic responses to climate change and human impact on the southeastern Tibetan Plateau during the past two centuries

Rapid population growth and economic development have led to increased anthropogenic pressures on the Tibetan Plateau, causing significant land cover changes with potentially severe ecological consequences. To assess whether or not these pressures are also affecting the remote montane‐boreal lakes on the SE Tibetan Plateau, fossil pollen and diatom data from two lakes were synthesized. The interplay of aquatic and terrestrial ecosystem response was explored in respect to climate variability and human activity over the past 200 years. Nonmetric multidimensional scaling and Procrustes rotation analysis were undertaken to determine whether pollen and diatom responses in each lake were similar and synchronous. Detrended canonical correspondence analysis was used to develop quantitative estimates of compositional species turnover. Despite instrumental evidence of significant climatic warming on the southeastern Plateau, the pollen and diatom records indicate very stable species composition throughout their profiles and show only very subtle responses to environmental changes over the past 200 years. The compositional species turnover (0.36–0.94 SD) is relatively low in comparison to the species reorganizations known from the periods during the mid‐ and early‐Holocene (0.64–1.61 SD) on the SE Plateau, and also in comparison to turnover rates of sediment records from climate‐sensitive regions in the circum arctic. Our results indicate that climatically induced ecological thresholds are not yet crossed, but that human activity has an increasing influence, particularly on the terrestrial ecosystem in our study area. Synergistic processes of post‐Little Ice Age warming, 20th century climate warming and extensive reforestations since the 19th century have initiated a change from natural oak‐pine forests to seminatural, likely less resilient pine‐oak forests. Further warming and anthropogenic disturbances would possibly exceed the ecological threshold of these ecosystems and lead to severe ecological consequences.     

The use of diatoms in monitoring the development of created wetlands at a sandmining site in Western Australia

Former sand-mining pits at Capel, 200 km south of Perth in Western Australia, have been rehabilitated into artificial wetlands since 1975–1979. A chain of fresh water lakes was created as a potential waterbird refuge and an area for passive recreation. Initially, the lakes had low pH, high ammonium, iron and manganese levels and low phosphorus concentration. The lakes were characterised by low diversity of diatoms dominated by acidophilous species. Following an increase in pH in the effluent water discharged into the lake from the mining process plant and landscaping of the lakes since 1988, the diversity of diatoms gradually increased. The system is now dominated by periphytic diatom communities, preferring high conductivity. There has been a marked transition in the diatom community from acidophilous to alkaliphilous species. Planktonic diatom blooms replaced dinoflagellate blooms. Concomitantly, there has been a dramatic increase in the diversity of invertebrates and waterbirds in these lakes. The value of diatoms in assessing the progressive development of created wetlands as self-sustaining ecosystems at sand mines in Australia is discussed.     

Evaluating diatom community composition in the absence of marked limnological gradients in the high Arctic: a surface sediment calibration set from Cornwallis Island (Nunavut,Canada)

Fossil diatom assemblages preserved within the sedimentary record in Arctic lakes provide the potential to reconstruct past changes in important limnological variables. During the summers of 1992 and 1993, we examined previously unstudied freshwater ecosystems on Cornwallis Island, Arctic Canada, with the specific objectives of (1) documenting the limnology and modern diatom assemblages from this region, and (2) determining which environmental variables most influence diatom species distributions. The Cornwallis Island study sites displayed the least amount of variance in measured water chemistry variables in comparison to nearly all of our labs’ previous freshwater surveys in the Arctic. The small limnological gradients precluded the development of a statistically robust diatom inference model, but perhaps more importantly, allowed us to explore variations in diatom composition in the absence of marked variations in water chemistry. Diatom species turnover was minimal, with the most common diatom taxa being Achnanthidium minutissima, Nitzschia perminuta, N. frustulum, with lesser percent abundances of Chaemaepinnularia soehrensis, Navicula chiarae, Psammothidium marginulata, and A. kryophila. A small number of study sites differed from the majority with respect to water chemistry (e.g., coastal sites with high specific conductivities) and habitat availability (e.g., ephemeral ponds with extensive moss habitats), and these sites had markedly different diatom assemblages. These data reinforce previous observations that water chemistry and other climate-related factors are the primary environmental controls influencing diatom distributions at high latitudes. Electronic supplementary material The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Sedimentary diatom concentrations and accumulation rates as predictors of lake trophic state

Diatom concentrations in surface sediments are positively correlated with limnetic chlorophyll a concentrations in Florida (USA) lakes. Using this relationship, I examine models that provide quantitative inferences for trophic state in historical applications.The best model predicts chlorophyll a trophic state index (TSI) values from log-transformed diatom concentrations and explains approximately half the variance in the dependent variable. Diatom accumulation rates are not better than sedimentary diatom concentrations as predictors of TSI. The entire diatom assemblage is as sensitive an indicator of TSI as are the planktonic diatoms alone. A model that considers the ecological preferences of specific taxa was found to be a better predictor than the model based on total diatom concentration.The sedimentary diatom concentration model provides a useful method for assessing historical changes in primary productivity, except in lakes where factors (e.g., silica limitation, blue-green bacterial inhibition) limited diatom production, or post-depositional changes removed sedimentary diatoms. TSI inferences are presented for sediment cores from two Florida lakes, one of which demonstrates a problematic application, and the other of which does not.