Combining limnological and palaeolimnological data to disentangle the effects of nutrient pollution and climate change on lake ecosystems: problems and potential

1. As long‐term observational lake records continue to lengthen, the historical overlap with lake sediment records grows, providing increasing opportunities for placing the contemporary ecological status of lakes in a temporal perspective. 2. Comparisons between long‐term data sets and sediment records, however, require lake sediments to be accurately dated and for sediment accumulation rates to be sufficiently rapid to allow precise matching with observational data. 3. The critical role of the sediment record in this context is its value in tracking the changing impact of human activity on a lake from a pre‐disturbance reference through to the present day. 4. Here, we use data from a range of lakes across Europe presented as case studies in this Special Section. The seven sites considered all possess both long‐term observational records and high‐quality sediment records. Our objective is to assess whether recent climate change is having an impact on their trophic status and in particular whether that impact can be disentangled from the changes associated with nutrient pollution. 5. The palaeo‐data show clear evidence for the beginning of nutrient pollution varying from the mid‐nineteenth century at Loch Leven to the early and middle twentieth century at other sites. The monitoring data show different degrees of recovery when judged against the palaeo‐reference. 6. The reason for limited recovery is attributed to continuing high nutrient concentrations related to an increase in diffuse nutrient loading or to internal P recycling, but there is some evidence that climate change may be playing a role in offsetting recovery at some sites. If this is the case, then lake ecosystems suffering from eutrophication may not necessarily return to their pre‐eutrophication reference status despite the measures that have been taken to reduce external nutrient loading. 7. The extent to which future warming might further limit such recovery can be evaluated only by continued monitoring combined with the use of palaeo‐records that set the pre‐eutrophication reference.     

Eutrophication,recovery and temperature in Lake Mjøsa: detecting trends with monitoring data and sediment records

1. How climate warming may interact with other pressures on aquatic ecosystems is an important issue for research and management. We combined lake monitoring data with a palaeolimnological study to explore the combined effects of eutrophication and subsequent oligotrophication with a long‐term temperature increase in epilimnetic waters. Our goals were (i) to evaluate how well sediment‐based reconstructions reflect the instrumental observations, (ii) to use the palaeo‐record to characterise a reference state for the lake and (iii) to explore whether data from the sediment record can aid in separating the effects of nutrient load and temperature in a large and deep lake. 2. Lake Mjøsa is a large and deep lake in south‐eastern Norway. Eutrophication symptoms peaked in the 1970s, which led to extensive measures to reduce the phosphorus load. A monitoring programme has run continuously from 1972. Monitoring has documented a marked decrease in phosphorus load and algal biomass and also revealed an increase in epilimnetic temperature and extended summer stratification. 3. Records of algal pigments and diatoms were extracted from sediment cores taken from 236 m depth. The pigment record documented dramatic changes in lake production consistent with the monitoring record. The diatom record reflected well the eutrophication history of the lake and also demonstrated that the assemblage of the recent recovery stage differs from that of the pre‐eutrophication period. 4. Ordination of diatom assemblages over time constrained by proxies for nutrient load and temperature indicated that the diatom assemblage correlated with both factors, which together accounted for 60% of the variation in diatom composition. No interaction was detected between these factors. The results suggest that the diatom assemblage has responded to varying nutrient loads as well as to changes in temperature and/or factors that correlate with temperature. 5. Reconstructions of algal biomass and total phosphorus content mirrored known changes through the monitoring period, although the absolute phosphorus estimates were too high relative to the instrumental record. The sediment record from Lake Mjøsa provides a baseline for lake production in terms of algal pigments and organic contents, and for the diatom assemblage composition in a pristine stage.     

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.     

Invertebrate food sources for waterbirds provided by the reconstructed wetland of Nyirkai-Hany,northwestern Hungary

Detailed studies of the macroinvertebrate benthos and zooplankton communities in Loch Leven, the largest shallow lowland lake in Scotland, UK, were carried out from 1966 to 1973 as part of the International Biological Programme (IBP). The results revealed a reduction in species diversity that was attributed to increasing eutrophication. This work provides a baseline against which the response of the invertebrate communities to subsequent changes in management can be assessed. This article compares macroinvertebrate benthos and zooplankton data from the IBP study with the post-IBP era during which changes at Loch Leven included a 60% reduction in the phosphorus input from external sources and variations in fish stocking rates. Only in recent years has there been evidence of ecological recovery by the invertebrate communities: the number of macroinvertebrate and zooplankton taxa has increased (including taxa considered to be sensitive to nutrient enrichment) and invertebrate abundances have declined. These changes appear to reflect the improvements in water quality and habitat conditions at Loch Leven that have occurred as a result of the recent reduction in nutrient loads, albeit with a substantial delay before any ecological response could be detected. This time lag in recovery has important implications for assessing improvements in the ecological status of other lake systems, as is required by the EU Water Framework Directive.     

Climate drivers of diatom distribution in shallow subarctic lakes

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A history of scientific research at Loch Leven,Kinross, Scotland

Loch Leven is a large, shallow lake in lowland Scotland, UK. Scientific research began here almost 200 years ago. Early research characterised the biodiversity and physical characteristics of the loch, providing an important historical background for future research. In the mid-1960s, this ad hoc approach was superseded by a more structured research programme under the umbrella of the International Biological Programme. This was the beginning of the Loch Leven long-term monitoring programme. Today, the results of these studies form one of the longest and most comprehensive limnological datasets for shallow freshwater lakes in the world, comprising more than 500 physical, chemical and biological variables collected at two-weekly intervals. To celebrate the 40th anniversary of the start of the long term monitoring programme, and to highlight the scientific investigations still being conducted at Loch Leven, the NERC Centre for Ecology & Hydrology (CEH) organised a symposium entitled “Loch Leven: 40 years of scientific research” in Kinross, Scotland, UK, on 11 December 2008. This examined the role of long-term monitoring in developing our understanding of the links between pollution, climate change and ecological responses in shallow lakes. This article introduces a series of papers summarising the scientific results presented at this meeting.     

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.     

Changes in aquatic macrophyte communities in Loch Leven: evidence of recovery from eutrophication?

This article assesses changes in the macrophyte community of Loch Leven over a period of 100 years. Evidence is presented that shows that these changes are associated with eutrophication and with subsequent recovery from eutrophication when anthropogenic nutrient inputs to the loch were reduced. This study uses macrophyte survey data from 1905, 1966, 1972, 1975, 1986, 1993, 1999 and 2008. In each of the four most modern surveys, the loch was divided into 19 sectors, each with at least one transect ranging from the shallowest to the deepest occurrence of macrophytes. From these data, a range of indicators of recovery were derived at the whole lake scale: the relative abundance of taxa, taxon richness and evenness. All of these metrics showed an improvement since 1972. Species richness, measured at the scales of survey sector and individual samples, also appeared to have increased in recent years. All of these measures, coupled with ordination of the presence/absence composition data from all survey years, indicate that the macrophyte community in the loch is recovering towards the state that was recorded in 1905.     

Climate forcing of diatom productivity in a lowland,eutrophic lake: White Lough revisited

1. In cultural landscapes, lake response to climate can be masked by land‐use change and nutrient loss from their catchments. Palaeolimnological methods were used to reconstruct the ecological response of diatoms in a eutrophic lowland lake (White Lough, Co. Tyrone, Northern Ireland) to altered nutrient P loading and precipitation variability over c. 100 years. 2. ²¹⁰Pb‐dated sediment cores were analysed to determine diatom assemblage variability, biogenic silica concentration, geochemical phosphorus concentration and accumulation rate. Manure P and agricultural N surplus data were collated from documentary sources. Long‐term trends in annual temperature and precipitation were derived from the Armagh Observatory. 3. Diatom community turnover from 1890 until c. 1960 was limited, and assemblages were dominated by Aulacoseira subarctica; after this date, changes primarily reflected a eutrophication sequence owing to increased diffuse nutrient inputs associated with intensification of land use (external P loading increased by a factor of three). 4. Diatom and biogenic Si profiles were compared with North Atlantic Oscillation (NAO) records, an index of regional weather patterns. Biogenic Si exhibited a c. 7‐year cycle, which tracked a cycle of similar timescale in the Armagh climate record for dry summers. In turn, this cycle was related to the variation in the NAO. 5. Monitoring data from 1971 to 2007 of nitrate exports from the Blackwater River showed that these too followed a roughly 7‐year cycle at least up to 2000, in which dry summers were followed by sharp increases in nitrate export. It is argued that diatom production in White Lough reflects the cyclic behaviour in nitrate loading and the constraints that nitrogen availability places on the spring diatom bloom in a lake that is dominated by cyanobacteria.     

Climate-lake interactions recorded in varved sediments from a Swedish boreal forest lake

A 1100-year long record of lake ecosystem response to climate and catchment change with precise chronological control is reported. Diatom and pollen assemblages of an annually laminated (varved) sediment from a northern Swedish lake (Kassjön, Våsterbotten) were used as records of lake diatom communities and catchment vegetation. These data were compared with summer temperature estimates based on tree-ring records of the same geographical area to identify the effects of climate change and catchment disturbance on diatom assemblages in the lake. In a canonical ordination, 23% of the variability in the total diatom assemblages for the period AD1040–1804 was accounted for by changes in pollen data which reflect agricultural development in the catchment. Diatom species richness, however, exhibited a stronger relationship with summer temperature and, significantly, declined with the lower temperatures associated with the Little Ice Age minimum (early 17th century). Summer temperature accounted for 23% of the variability in diatom species richness 20 years later. The mechanism behind this time-lag is unclear, but may be related to catchment-mediated effects, given recent evidence for lags in the response of boreal-forest vegetation regeneration cycles to climatic variability. These results suggest that climate-related effects on lakes occurring over medium timescales can be resolved in lake sediments. Moreover, it is possible to identify these effects despite cultural-related signals, but as the latter become more extreme in the late 20th century the climate signal is obscured.     

Major changes in trophic dynamics in large, deep sub-alpine Lake Maggiore from 1940s to 2002: a high resolution comparative palaeo-neolimnological study

1. We describe the changes in trophic dynamics in Lake Maggiore from c. 1943 to 2002 using subfossil cladoceran data from a high resolution sediment record, long‐term contemporary data series and historical information. During this period the lake went through a eutrophication phase until 1980 followed by oligotrophication. 2. During the eutrophication period a major increase occurred in the abundance of Chydorus sphaericus, the proportion of planktonic cladocerans and total abundance of cladocerans in the sediment. Since 1980 the abundance declined again and subfossil Eubosmina mucro length and contemporary Daphnia body length increased, most probably as a result of higher abundance of invertebrate predators. 3. Changes in the fish stock composition caused by the introduction of exotic fish during the pre‐eutrophication period and a complete ban on fishing because of Dichloro‐diphenil‐ethanes (DDTs) pollution of the lake (during oligotrophication) could also be detected in the community assemblage and size structure of the sediment zooplankton. 4. We found good correspondence between trophic changes inferred from cladoceran subfossils (community composition, size and predation pressure) and contemporary data, suggesting that sediment samples can be used to infer past development in trophic dynamics, including predation by fish and pelagic invertebrates in lakes with scarce neolimnological data. 5. Furthermore, by combining palaeolimnological cladoceran data rarely obtained from contemporary samples (e.g. benthic and plant‐associated cladocerans, mucro length of bosminids) with contemporary data of organisms poorly represented in the sediment record (e.g. remains of Bythotrephes and fishes) a more complete understanding of changes in trophic dynamics was obtained. 6. The detection in the sediments of meteorological events whose effects on zooplankton had been recorded in the long‐term studies also provided evidence that eutrophication tends to override climate signals. 7. We conclude that a combined palaeo‐neolimnological approach can be a powerful tool for elucidating past changes in the trophic dynamics of lakes and the interaction with climate induced changes, not least when high resolution sediment records are available.     

Combining monitoring,models and palaeolimnology to assess ecosystem response to environmental change at monthly to millennial timescales: the stability of Blue Lake,North Stradbroke Island,Australia

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Asynchronous onset of eutrophication among shallow prairie lakes of the Northern Great Plains,Alberta, Canada

Coherent timing of agricultural expansion, fertilizer application, atmospheric nutrient deposition, and accelerated global warming is expected to promote synchronous fertilization of regional surface waters and coherent development of algal blooms and lake eutrophication. While broad‐scale cyanobacterial expansion is evident in global meta‐analyses, little is known of whether lakes in discrete catchments within a common lake district also exhibit coherent water quality degradation through anthropogenic forcing. Consequently, the primary goal of this study was to determine whether agricultural development since ca. 1900, accelerated use of fertilizer since 1960, atmospheric deposition of reactive N, or regional climate warming has resulted in coherent patterns of eutrophication of surface waters in southern Alberta, Canada. Unexpectedly, analysis of sedimentary pigments as an index of changes in total algal abundance since ca. 1850 revealed that while total algal abundance (as β‐carotene, pheophytin a) increased in nine of 10 lakes over 150 years, the onset of eutrophication varied by a century and was asynchronous across basins. Similarly, analysis of temporal sequences with least‐squares regression revealed that the relative abundance of cyanobacteria (echinenone) either decreased or did not change significantly in eight of the lakes since ca. 1850, whereas purple sulfur bacteria (as okenone) increased significantly in seven study sites. These patterns are consistent with the catchment filter hypothesis, which posits that lakes exhibit unique responses to common forcing associated with the influx of mass as water, nutrients, or particles.     

The sensitivity of phytoplankton in Loch Leven (U.K.) to changes in nutrient load and water temperature

1. Loch Leven is a shallow, eutrophic lake in Scotland, U.K. It has experienced much change over the 30 years that it has been studied; this has primarily been due to reduced nutrient loads to the lake through active catchment management. Its recovery has been slow and, therefore, we used a phytoplankton community model (PROTECH) to test its sensitivity to changing nutrient loads and water temperature.
2. PROTECH was initialized to simulate the observed phytoplankton community in 1995 and was then repeatedly run through a combination of step-wise changes in water temperature and nutrient load (two treatments were simulated for nutrient load: one changing both nitrate and phosphorus, and one changing just phosphorus). The effect on total chlorophyll- a concentration, cyanobacteria abundance and phytoplankton diversity was examined.
3. Whilst changes in temperature had little effect, variations in the nutrient load produced a range of responses. Increasing only the phosphorus load caused a large increase in Anabaena abundance and total chlorophyll- a concentration. However, the opposite response was recorded when nitrate load was changed as well, with Anabaena increasing its biomass under reduced nutrient load scenarios.
4. The key factor determining the type of response appeared to be nitrogen availability. Anabaena , a nitrogen fixer, could exploit the phosphorus resource of Loch Leven under limiting nitrogen conditions, allowing it to dominate under most of the scenarios tested apart from those supplying extra nitrogen to the lake. The model predictions agree with the observed data, which show that Anabaena continues to dominate the summer phytoplankton bloom in Loch Leven despite the considerable reduction in phosphorus supply from the catchment. This research provides a possible explanation for this.     

星云湖硅藻群落响应近现代人类活动与气候变化的过程

随着人类活动的增强与全球气候变暖的持续,近年来云南湖泊的生态系统功能持续退化,而目前对云南湖泊生态系统的研究还主要集中于单一环境压力的生态效应。以星云湖为研究对象,通过沉积物记录与现代监测资料,识别在湖泊富营养化、气候变化以及人类强烈干扰下硅藻群落结构响应的过程,并甄别驱动群落变化的主要环境压力及其强度。结果显示随着湖泊生产力水平(如沉积物叶绿素a浓度)的增加,硅藻物种组成发生了明显的变化,主成分分析表明了水体富营养化是驱动群落变化的主要环境因子(r=-0.63,P0.001)。简约模型与方差分解的结果表明近200年来(钻孔长度38cm),湖泊营养水平和水动力是驱动星云湖硅藻群落变化的主要环境因子,分别解释了群落变化的18.8%和2.9%;而1951年以后,湖泊营养水平和温度分别解释了硅藻群落结构变化的31.4%和26.8%。研究结果表明了硅藻群落长期变化的主控因子是湖泊营养水平,而人类活动及气候变化等可以通过改变湖泊水动力及湖水温度来驱动硅藻群落的演替,同时抚仙湖-星云湖的连通性也对硅藻群落的演替产生了一定影响。     

Maximum growing depth of macrophytes in Loch Leven,Scotland, United Kingdom,in relation to historical changes in estimated phosphorus loading

Eutrophication is common in shallow lakes in lowland areas. In their natural state, most shallow lakes would have clear water and a thriving aquatic plant community. However, eutrophication often causes turbid water, high algal productivity, and low species diversity and abundance of submerged macrophytes. A key indicator of the ecological state of lake ecosystems is the maximum growing depth (MGD) of aquatic plants. However, few studies have yet quantified the relationship between changes in external phosphorus (P) input to a lake and associated variation in MGD. This study examines the relationship between these variables in Loch Leven, a shallow, eutrophic loch in Scotland, UK. A baseline MGD value from 1905 and a series of more recent MGD values collected between 1972 and 2006 are compared with estimated P loads over a period of eutrophication and recovery. The results suggest a close relationship between changes in MGD of macrophytes and changes in the external P load to the loch. Variation in MGD reflected the ‘light history’ that submerged macrophytes had been exposed to over the 5-year period prior to sampling, rather than responding to short term, within year, variations in water clarity. This suggests that changes in macrophyte MGD may be a good indicator of overall, long term, changes in water quality that occur during the eutrophication and restoration of shallow lakes.     

The diatom assemblage of a marl core from Linton Loch

Summary

Diatom data from Linton Loch show five diatom assemblage zones, together with stratigraphic evidence indicating some aspects of the development of the lake basin during the Flandrian. Results show an early Flandrian dominance of planktonic diatom species in the lake waters with the later development of epiphytic species as the hydrosere encroached. Following deforestation in the catchment, water levels rose in the basin when benthic diatoms became dominant.     

Centennial‐scale changes to the aquatic vegetation structure of a shallow eutrophic lake and implications for restoration

1. We investigate long‐term (>200 years) changes to the composition and spatial structure of macrophyte communities in a shallow, eutrophic lake (Barton Broad, eastern England) and consider the implications for lake restoration. 2. Historical macrophyte data were assembled from a variety of sources: existing plant databases, museum herbaria, journal articles, old photographs and eyewitness accounts. Additionally, two types of sediment core sample were analysed for plant macro‐remains and pollen; bulk basal samples from multiple core sites analysed to provide information on ‘pre‐disturbance’ macrophyte communities and two whole cores analysed to determine historical change. 3. Prior to the late 1800s, macrophyte communities were diverse and included a multilayered mosaic of short‐stature submerged taxa and taller submerged and floating‐leaved species. With the progression of eutrophication after around 1900, the former community was displaced by the latter. Diversity was maintained, however, since an encroaching Schoenoplectus–nymphaeid swamp generated extensive patches of low‐energy habitat affording refugia for several macrophytes otherwise unable to withstand the hydraulic forces associated with open water conditions. When this swamp vegetation disappeared in the 1950s, many of the ‘dependent’ aquatic macrophytes also declined leaving behind a sparse, species‐poor community (as today) resilient to both eutrophication and turbulent open waters. 4. The combination of historical and palaeolimnological data sources offers considerable benefits for reconstructing past changes to the aquatic vegetation of lakes and for setting restoration goals. In this respect, our study suggests that successful restoration might often be better judged by reinstatement of the characteristic structure of plant communities than the fine detail of species lists; when nutrients are low and the structure is right, the right species will follow.     

The recent diatom history of Loch Leven, Kinross

Diatoms in the top metre of lake sediment have been studied for indications or recent changes in the environment of Loch Leven. The diatoms indicate a eutrophic, alkaline lake with Melosira italica ssp. subarctica, Tabellaria Flocculosa and Stephanodiscus astraea the dominant taxa at the base of the 1-m section. There is then an appeance of Melosira ambigua and Stephanodiscus hantzschii suggesting a richer environment. Post A.D. 1816 changes in taxa suggest possible correlation with the lowering of lake level in 1830 and the dominance of Diatoma elongatum, Cyclotella pseudostelligera, Stephanodiscus astraea var. minutula, S. tenuis, S. hantzschii, and Synedra rumpens and var. soctica in zone 4 agrees with present-day plankton records.     

The relationship between Trichocerca pusilla (Jennings), Aulacoseira spp. and water temperature in Loch Leven,Scotland, U.K.

Loch Leven is a shallow, eutrophic lake in the Scottish lowlands that is famous for its brown trout (Salmo trutta L.) fishery. Studies of planktonic rotifer populations began here in January 1977. Since then, samples have been collected and analysed at more or less weekly intervals. Additional information on the composition and abundance of phytoplankton and crustacean zooplankton species, and on a variety of physical and chemical determinants, has been recorded on each sampling occasion.Long-term datasets, such as that described above, are invaluable for identifying interactions between components of the plankton that only appear for short periods each year, as these interactions would probably be overlooked in data spanning a shorter period of time. This study uses the long-term data from Loch Leven to examine the food and temperature requirements of the summer rotifer species Trichocerca pusilla (Lauterborn). The results suggest that T. pusilla prefers water temperatures above 12 °C and that it feeds, primarily, on the filamentous diatom Aulacoseira spp. During the summer months, its abundance was closely related to the availability of this diatom. When filaments of Aulacoseira spp. were abundant, rotifer densities reached 1000–3000 ind. l^–1 and when they were scarce (e.g. 1980, 1997 and 1998) T. pusilla densities also remained low (i.e. less than 100 ind. l^–1). The reason for the success or failure of Aulacoseira during the summer months each year is unclear but, in general, its abundance was related to the availability of dissolved silica in the water.