A 250 year comparison of historical, macrofossil and pollen records of aquatic plants in a shallow lake

1. Sedimentary remains of aquatic plants, both vegetative (turions, leaves, spines) and reproductive (fruits, seeds, pollen), may provide a record of temporal changes in the submerged vegetation of lakes. An independent assessment of the degree to which these remains reflect past floristic change is, however, rarely possible. 2. By exploiting an extensive series of historical plant records for a small shallow lake we compare plant macrofossil (three cores) and pollen (one core) profiles with the documented sequence of submerged vegetation change since c. 1750 AD. The data set is based on 146 site visits with 658 observations including 42 taxa classified as aquatic, spanning 250 years. 3. Approximately 40% of the historically recorded aquatic taxa were represented by macro‐remains. In general macrofossils underestimated past species diversity, with pondweeds (three of eight historically recorded Potamogeton species were found) particularly poorly represented. Nonetheless, several taxa not reported from historical surveys (e.g. Myriophyllum alterniflorum and Characeae) were present in the sediment record. 4. The pollen record revealed taxa which left no macro‐remains (e.g. Littorella uniflora), and the macrofossil record provided improved taxonomic resolution for some taxa (e.g. Potamogeton) and a more reliable record of persistence, appearance and loss of others (e.g. Myriophyllum spp. and Nymphaeaceae). 5. Detrended correspondence analysis indicated that changes in the community composition evidenced by the palaeolimnological and historical records were synchronous and of a similar magnitude. Both records pointed to a major change at around 1800, with the historical record suggesting a more abrupt change than the sedimentary data. There was good agreement on a subsequent change c. 1930. 6. The palaeolimnological data did not provide a complete inventory of historically recorded species. Nevertheless, these results suggest that combined macrofossil and pollen records provide a reliable indication of temporal change in the dominant components of the submerged and floating‐leaved aquatic vegetation of shallow lakes. As such palaeolimnology may provide a useful tool for establishing community dynamics and successions of plants over decadal to centennial timescales.     

Micro‐scale structure in the chemistry and biology of a shallow lake

1. Repeat sampling in daytime within a lily (Nuphar lutea) bed and in open water showed distinct heterogeneities in the three‐dimensional distributions of water chemistry and planktonic organisms on centimetre to decimetre scales. 2. Vertical gradients of physico‐chemical variables that did not exist at dawn developed during the day in both sites, as available nutrients were released from the sediments and were consumed towards the surface. 3. Distributions of algal standing crop suggest limitation by both nutrients and grazing. 4. Marked variability in distributions may question the assumptions often made about the homogeneity of plankton and available nutrient distributions in open water and in macrophyte stands of shallow lakes. Although simple sampling regimes for monitoring of water quality may be adequate for many purposes, they miss a fine structure in the water that is inherently interesting in understanding the underlying processes of plankton function.     

Long‐term dynamics of submerged macrophytes and algae in a small and shallow,eutrophic lake: implications for the stability of macrophyte‐dominance

1. Submerged macrophyte and phytoplankton components of eutrophic, shallow lakes have frequently undergone dynamic changes in composition and abundance with important consequences for lake functioning and stability. However, because of a paucity of long‐term survey data, we know little regarding the nature, direction and sequencing of such changes over decadal–centennial or longer timescales. 2. To circumvent this problem, we analysed multiple (n = 5) chronologically correlated sediment cores for plant macro‐remains and a single core for pollen and diatoms from one small, shallow, English lake (Felbrigg Hall Lake, Norfolk, U.K.), documenting 250 years of change to macrophyte and algal communities. 3. All five cores showed broadly similar stratigraphic changes in macrophyte remains with three distinct phases of macrophyte development: Myriophyllum–Chara–Potamogeton (c. pre‐1900), to Ceratophyllum–Chara–Potamogeton (c. 1900–1960) and finally to Zannichellia–Potamogeton (c. post‐1960). Macrophyte species richness declined from at least 10 species pre‐1900 to just four species at the present day. Additionally, in the final Zannichellia–Potamogeton phase, a directional shift between epi‐benthic and phytoplankton‐based primary production was indicated by the diatom data. 4. Based on macrophyte–seasonality relationships established for the region, concomitant with the final shift to Zannichellia–Potamogeton, we infer a reduction in the seasonal duration of plant dominance (plant‐covered period). Furthermore, we hypothesise that this change in species composition resulted in a situation whereby macrophyte populations were seasonally ‘sandwiched’ between two phytoplankton peaks in spring and late summer as observed in the contemporary lake. 5. We suggest that eutrophication‐induced reductions in macrophyte species richness, especially if the number of plant‐seasonal strategies is reduced, may constrict the plant growing season. In turn, this may render a shallow lake increasingly vulnerable to seasonal invasions of phytoplankton resulting in further species losses in the plant community. Thus, as part of a slow (over perhaps 10–100s of years) and self‐perpetuating process, macrophytes may be gradually pushed out by phytoplankton without the need for a perturbation as required in the alternative stable states model of plant loss.     

Clear,crashing, turbid and back – long‐term changes in macrophyte assemblages in a shallow lake

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Seasonal water quality of shallow and eutrophic Lake Pamvotis, Greece: implications for restoration

Lake Pamvotis is a moderately sized (22 km²) shallow (z _avg=4 m) lake with a polymictic stratification regime located in northwest Greece. The lake has undergone cultural eutrophication over the past 40 years and is currently eutrophic (annual averages of FRP=0.07 mg P l^-1, TP=0.11 mg P l^-1, NH₄ ⁺=0.25 mg N l^-1, NO₃ ^–=0.56 mg N l^-1). FRP and NH₄ ⁺ levels are correlated to external loading from streams during the winter and spring, and to internal loading during multi-day periods of summer stratification. Algal blooms occurred in summer (July–August green algae, August–September blue-green algae), autumn (October blue-green algae and diatoms), and winter (February diatoms), but not in the spring (March–June). The phytoplankton underwent brief periods of N- and P-limitation, though persistent low transparency (secchi depth of 60–80 cm) also suggests periods of light limitation. Rotifers counts were highest from mid-summer to early autumn whereas copepods were high in the spring and cladocerans were low in the summer. Removal of industrial and sewage point sources a decade ago resulted in a decrease in FRP. A phosphorus mass balance identified further reductions in external loading from the predominately agricultural catchment will decrease FRP levels further. The commercial fishery and lake hatchery also provides opportunities to control algal biomass through biomanipulation measures.     

The development and persistence of alternative ecosystem states in a large,shallow lake

1. Palaeolimnological data were used to investigate drivers of the community of primary producers in Lake Mattamuskeet, North Carolina, U.S.A. This is a large, shallow lake with two basins currently dominated by phytoplankton and macrophytes. The two basins were divided in 1940 by the building of a roadway across the lake, which also corresponded with the divergence in their ecosystem state. 2. Photosynthetic pigments, organic matter and nutrients (P, N, C, S) were analysed in sediment cores from each basin to reconstruct the primary producer community over the past c. 100 years. We sought to answer two questions. First, what changes to the ecosystem resulting from the building of the roadway caused the development of different primary producer communities in the two basins? Second, why have the alternative ecosystem states persisted despite a variety of human perturbations since 1940? 3. K‐means cluster analysis and principal component analysis were applied to identify three sediment types based on photosynthetic pigment data: sediments indicating low productivity (low pigment concentrations), sediments associated with macrophytes (chlorophyll a and b) and with phytoplankton (alloxanthin and aphanizophyll). In addition, other palaeolimnological proxies measured, such as loss on ignition, total phosphorus, total organic carbon/total nitrogen and other nutrients, were different in post‐1940 sediments within the two basins. 4. These differences suggest characteristics, such as nutrient cycling, water depth and other physical changes resulting from roadway construction, combined to establish and maintain the differing communities of primary producers in the two basins. Furthermore, Fe/S dynamics and waterfowl herbivory probably contributed to the development of the two ecosystem states.     

Natural and anthropogenic forcing of aquatic macrophyte development in a shallow Danish lake during the last 7000 years

Aim To investigate the long‐term changes in aquatic vegetation in a lowland, shallow lake, and to assess the relationship between aquatic vegetation and natural and anthropogenic catchment changes. Location Gundsømagle Sø, Zealand, Denmark: a shallow (mean depth 1.2 m), hypereutrophic lake (mean annual total phosphorus (TP) c. 700 μg TP L^?1) located in a predominantly agricultural catchment (88% cultivated land). The lake is presently devoid of macrophytes. Methods One hundred and forty‐seven contiguous samples from a sediment core (taken in 2000) were analysed for macrofossil remains together with loss‐on‐ignition and dry weight. From an earlier sediment core (taken in 1992), 67 samples were analysed for pollen and the two cores were correlated using the ignition residue profiles. Core chronology was determined by ²¹⁰Pb and ¹³⁷Cs dating of the recent lake sediments, while older sediments were dated by pollen‐stratigraphical correlation, as ¹⁴C dating proved problematical. Aquatic macrofossil abundance was used to reconstruct past changes in the lake's plant community and water‐level. The contemporary catchment land‐use change was inferred from sedimentary pollen data, and soil erosion to the lake was deduced from the minerogenic content of the lake sediments. Results The macrofossil record covers the last 7000 years, but aquatic plant remains were scarce prior to c. 1300 bc . After this date the abundance of submerged and emergent macrophyte remains increased dramatically, paralleled by an increase in sediment minerogenic matter and non‐arboreal pollen (NAP). Aquatic plant remains were abundant for more than 3000 years until the mid 1900s. Macrofossils of Linum usitatissimum (L.) (flax) and high pollen percentages of ‘Cannabis type’ (hemp) were recorded in periods between c. 1150 bc and 1800 ad . Main conclusions Our study suggests that, between c. 5000 bc and 1300 bc , the submerged plant community was confined to the littoral zone. From 1300 bc onwards, the submerged macrophyte vegetation expanded rapidly across the lake bed, presumably as a response to lake shallowing caused by a combination of climatic‐induced water‐level lowering and enhanced erosional infilling of the lake basin due to intensified anthropogenic activities in the catchment. The lake was meso‐eutrophic and had an extensive and diverse aquatic flora for more than 3000 years, until the middle of the twentieth century. In periods between c. 1150 bc and 1800 ad , the lake experienced direct anthropogenic impact from retting of fibre plants (Linum and Cannabis). Over the last 200 years, erosional infilling of the lake basin increased drastically, probably as a result of agricultural intensification. In the twentieth century, the lake was strongly affected by nutrient enrichment from both point sources (sewage from built‐up areas) and diffuse agricultural run‐off which led to hypertrophic conditions and the collapse of the submerged vegetation c. 1950–60. The concept of ‘naturalness’ and the implications for lake conservation are discussed.     

Combining contemporary ecology and palaeolimnology to understand shallow lake ecosystem change

1. Palaeolimnology and contemporary ecology are complementary disciplines but are rarely combined. By reviewing the literature and using a case study, we show how linking the timescales of these approaches affords a powerful means of understanding ecological change in shallow lakes. 2. Recently, palaeolimnology has largely been pre‐occupied with developing transfer functions which use surface sediment‐lake environment datasets to reconstruct a single environmental variable. Such models ignore complex controls over biological structure and can be prone to considerable error in prediction. Furthermore, by reducing species assemblage data to a series of numbers, transfer functions neglect valuable ecological information on species’ seasonality, habitat structure and food web interactions. These elements can be readily extracted from palaeolimnological data with the interpretive assistance of contemporary experiments and surveys. For example, for one shallow lake, we show how it is possible to infer long‐term seasonality change from plant macrofossil and fossil diatom data with the assistance of seasonal datasets on macrophyte and algal dynamics. 3. On the other hand, theories on shallow lake functioning have generally been developed from short‐term (<1–15 years) studies as opposed to palaeo‐data that cover the actual timescales (decades–centuries) of shallow lake response to stressors such as eutrophication and climate change. Palaeolimnological techniques can track long‐term dynamics in lakes whilst smoothing out short‐term variability and thus provide a unique and important means of not only developing ecological theories, but of testing them. 4. By combining contemporary ecology and palaeolimnology, it should be possible to gain a fuller understanding of changing ecological patterns and processes in shallow lakes on multiple timescales.     

Chironomid stratigraphy in the shallow and eutrophic Lake Søbygaard, Denmark: chironomid–macrophyte co-occurrence

1. A sediment core from the shallow, hypertrophic Lake Søbygaard (mean depth ∼1 m; [TP] 310 μg P L⁻¹) was analysed for subfossil remains to reconstruct chironomid community changes in relation to the succession and disappearance of aquatic macrophytes. 2. Species composition in the 1.10 m core indicates a succession from a 'naturally' eutrophic state to a hypertrophic state during recent centuries. Radiometric dating (²¹⁰Pb) of the uppermost 20 cm of the sediment core (∼1932–93) indicates that sediment accumulation rate had doubled in recent decades. 3. Changes in chironomid assemblages were in close agreement with changes in both diatoms and macrophyte remains in the same core. Distinct changes in chironomid communities reflect the eutrophication process and macrophyte succession through Chara , Ceratophyllum and Potamogeton dominance to the present state, with complete loss of submerged vegetation and dominance by phytoplankton. 4. The co-occurrence and relationship between aquatic macrophyte diversity and recent subfossil chironomid assemblages were assessed from an additional 25 Danish lakes. There was good agreement between the macrophyte and chironomid-based lake groupings. Overall, a significant difference ( P <0.001) was found in chironomid assemblages among lakes in different macrophyte classes. In a pair-wise comparison, the poorly buffered mesotrophic lakes and the alkaline eutrophic lakes had significantly different chironomid assemblages. 5. Chironomid taxa commonly reported to be associated with macrophytes ( Cricotopus , Endochironomus and Glyptotendipes ) were shown also to be indicators of highly productive lakes lacking abundant submerged vegetation.     

The eutrophication of shallow coastal lakes in Southwest England — understanding and recommendations for restoration,based on palaeolimnology,historical records,and the modelling of changing phosphorus loads

Palaeolimnological studies of sediments from Slapton Ley and Loe Pool, two coastal freshwater lakes in Southwest England, show that in the period since 1945, they have been eutrophicated by nutrient inputs from intensification of agriculture, but also from sewage effluent. Two simple models have been used to identify the main sources of catchment outputs, and in the case of Slapton Ley, to evaluate historical changes in land use, and their likely effect on lake trophic status.Restoration strategies may also be evaluated using the same models. They suggest that in order to reduce loads on either lake to within OECD permissible limits, not only will all sewage inputs need to be prevented, and non-phosphate detergents used, but also losses from agricultural land must be reduced. This could take the form of the keeping of fewer cattle (the main source of organic nitrogen and phosphorus in both catchments), or the zoning of the respective catchments so that steep slopes close to riparian zones are not used, as at present, for the grazing of livestock.A better option, however, would appear to be the establishment along most of the rivers draining into these lakes, of buffer strips of woodland at least 15 m wide. According to the models, this measure, along with treatment or diversion of sewage effluent, would reduce phosphorus loads upon the lakes to within acceptable limits.     

Ecological influences on larval chironomid communities in shallow lakes: implications for palaeolimnological interpretations

1. To correctly interpret chironomid faunas for palaeoenvironmental reconstruction, it is essential that we improve our understanding of the relative influence of ecosystem variables, biotic as well as physicochemical, on chironomid larvae. To address this, we analysed the surface sediments from 39 shallow lakes (29 Norfolk, U.K., 10 Denmark) for chironomid head capsules, and 70 chironomid taxa (including Chaoborus) were identified. 2. The shallow lakes were selected over large environmental gradients of aquatic macrophytes, total phosphorus (TP) and fish communities. Redundancy analysis (RDA) identified two significant variables that explained chironomid distribution: macrophyte species richness (P < 0.001) and TP (P < 0.005). Generalised linear models (GLM) identified specific taxa that had significant relationships with both these variables. Macrophyte percentage volume infested (PVI) and species richness were significant in classifying the lake types based on chironomid communities under twinspan analysis, although other factors, notably nutrient concentrations and fish communities, were also important, illustrating the complexities of classifying shallow lake ecosystems. Lakes with plant species richness >10 all had relatively diverse (Hill’s N2) chironomid assemblages, and lakes with Hill’s N2 >10 all had TP <250 μg L⁻¹ and total fish densities <2 fish per m². 3. Plant density (PVI), and perhaps more importantly species richness, were primary controls on the distribution of chironomid communities within these lakes. This clearly has implications for palaeoenvironmental reconstructions using zoobenthos remains (i.e. chironomids) and suggests that they could be used to track changes in benthic/pelagic production and could be used as indicators of changing macrophyte habitat. 4. Measuring key biological gradients, in addition to physicochemical gradients, allowed the major controls on chironomid distribution to be assessed more directly, in terms of plant substrate, food availability, competition and predation pressure, rather than implying indirect mechanisms through relationships with nutrients. Many of these variables, notably macrophyte abundance and species richness, are not routinely measured in such studies, despite their importance in determining zoobenthos in temperate shallow lakes. 5. When physical, chemical and ecological gradients are considered, as is often the case with palaeo‐reconstructions rather than training sets chosen to maximise one gradient, complex relationships exist, and attempting to reconstruct a single trophic variable quantitatively may not be appropriate or reliable.     

Submerged aquatic vegetation correlations with depth and light attenuating materials in a shallow subtropical lake

A 3-year study was done to quantify the biomass of submerged aquatic vegetation (SAV) and its relationship with environmental attributes in Lake Okeechobee, the largest lake in the southeastern United States. Plants were sampled on 21 occasions at sites located along 15 fixed transects around the shoreline, giving rise to 721 observations of SAV species (Chara spp., Vallisneria americana, Hydrilla verticillata, Potamogeton illinoinensis) dry weight biomass. Environmental sampling focused on factors that attenuate light, including phytoplankton chlorophyll a (chl a), total suspended solids (TSS), non-volatile suspended solids (NVSS) and color. Depth and Secchi transparency also were measured. Based on regression analysis, NVSS was considerably more important in attenuating light than chl a or color. Total biomass of SAV varied from 0 to 271 g dw m^–2, with a mean of 4.7 g dw m^–2, and strong dominance by Chara. The SAV biomass was lower than average for Florida lakes, and may reflect the influence of suspended solids on underwater irradiance, as well as high water level in the late 1990s. Dense SAV was found only where depth was < 2 m and TSS < 20–30 mg l^–1. At locations where high biomass of SAV occurred, the plants may have influenced water quality, because concentrations of TSS, NVSS, and chl a were 2–3 fold lower than at sites with no plants. The potential effects of SAV also were apparent at a regional scale. The shoreline region of the lake displayed a pattern of rising and falling chl a and NVSS with water depth. This occurred both at sites with and without plants, suggesting that it may be driven by physical processes, such as water circulation patterns, which are influenced by depth. However, the pattern was dampened at sites with SAV, indicating a potential to influence these attributes of water quality.     

Seasonal changes in sediment and water chemistry of a subtropical shallow eutrophic lake

A field study was conducted (May 1981 to June 1982) to develop a data-base on seasonal changes of water and sediment chemistry of Lake Monroe (4 000 ha surface and ca. 2 m deep) located in central Florida, USA. This shallow eutrophic lake is a part of the St. Johns River. Quantitative samples of lake water and sediments were collected on a monthly basis from 16 stations and analyzed for various physico-chemical parameters. Relatively high levels of dissolved solids (mean electrical conductivity (EC) = 1832 µS cm¹) prevailed in the lake water, and seasonal changes in EC were probably associated with hydrologic flushing from external sources, such as incoming water from upstream as well as precipitation. Average monthly levels of total N and P during the study period were 1.82 and 0.21 mg l^–1, respectively. Nutrient concentrations in the water did not show any strong seasonal trends. Organic matter content of lake sediments ranged from 1 to 182 g C kg^–1 of dry sediment, reflecting considerable spatial variability. All nutrient elements in the sediments showed highly significant (P < 0.01) correlations with sediment organic C, though little or no significant relationship appeared at any sampling period between water and sediment chemistry of the lake. Temporal trends in water and sediment chemical parameters may have been concealed by periodic hydrologic flushing of the St. Johns River into Lake Monroe.Florida Agricultural Experiment Stations Journal Series No. 7836.     

Biodiversity changes in a shallow lake ecosystem: a multi-proxy palaeolimnological analysis

Biodiversity is a key measure of environmental quality in lake ecosystems. Lake biodiversity can be assessed using modern survey data, but typically these data only provide a ‘snap-shot’ measure and in most cases it is not possible to reconstruct temporal trends in biodiversity, so that human impacts can be detected. Palaeoecological techniques offer an alternative means of identifying changes in biodiversity over the period of historical records and far beyond, but there are problems associated with this approach. This is because only a select set of organisms leave a trace in the sediment record such that it is not usually possible to make reliable assessments of diversity changes within an entire taxonomic order (e.g. the algae). Moreover these organisms are typically from the lower levels of the trophic hierarchy (i.e. plants and insects). The problems of identifying changes in biodiversity from the palaeolimnological record are addressed with reference to Groby Pool, a shallow, eutrophic, medieval lake in the English Midlands, which has been subjected to eutrophication over the last 150 years. ²¹⁰Pb and ¹³⁷Cs-dated sediment cores have been used to estimate short-term alterations in the composition and diversity of three groups of indicators, representing different levels in the trophic cascade, namely diatoms, aquatic pollen and chironomids. By exploring relationships, both between these indicators and with archival macrophyte records, an assessment is made of eutrophication-related changes in overall habitat diversity at the ecosystem level. These data suggest that the lake has undergone considerable nutrient enrichment, resulting in the loss of a diverse, mesotrophic macrophyte flora from at least the turn of the century onwards and its replacement by a few highly competitive species tolerant of high nutrient concentrations. Reductions in macrophyte diversity seem to be reflected palaeoecologically by a decline in the diversity of fossil chironomid assemblages, related to the breakdown of particular host-plant relationships amongst the phytophagic species. However, diatom assemblages generally exhibit the opposite trend, which may be related to increases in macrophyte cover and increasing opportunities for the colonization of diverse epiphyte communities. The different fossil indicators have different limitations and merits, and for this reason a ‘multi-proxy’ approach is essential if meaningful inferences are to be made of changes in lake biodiversity using palaeoecological data.     

Heat‐wave effects on greenhouse gas emissions from shallow lake mesocosms

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Fish manipulation as a lake restoration tool in shallow,eutrophic, temperate lakes 2: threshold levels,long-term stability and conclusions

In order to evaluate short-term and long-term effects of fish manipulation in shallow, eutrophic lakes, empirical studies on relationships between lake water concentration of total phosphorus (P) and the occurrence of phytoplankton, submerged macrophytes and fish in Danish lakes are combined with results from three whole-lake fish manipulation experiments. After removal of less than 80 per cent of the planktivorous fish stock a short-term trophic cascade was obtained in the nutrient regimes, where large cyanobacteria were not strongly dominant and persistent. In shallow Danish lakes cyanobacteria were the most often dominating phytoplankton class in the P-range between 200 and 1 000μg P l⁻¹. Long-term effects are suggested to be closely related to the ability of the lake to establish a permanent and wide distribution of submerged macrophytes and to create self-perpetuating increases in the ratio of piscivorous to planktivorous fish. The maximum depth at which submerged macrophytes occurred, decreased exponentially with increasing P concentration. Submerged macrophytes were absent in lakes>10 ha and with P levels above 250–300μg P l⁻¹, but still abundant in some lakes<3 ha at 650μg P l⁻¹. Lakes with high cover of submerged macrophytes showed higher transparencies than lakes with low cover aboveca. 50μg P l⁻¹. These results support the alternative stable state hypothesis (clear or turbid water stages). Planktivorous fish>10 cm numerically contributed more than 80 per cent of the total planktivorous and piscivorous fish (>10 cm) in the pelagical of lakes with concentrations above 100μg P l⁻¹. Below this threshold level the proportion of planktivores decreased markedly toca. 50 per cent at 22μg P l⁻¹. The extent of the shift in depth colonization of submerged macrophytes and fish stock composition in the three whole-lake fish manipulations follows closely the predictions from the relationships derived from the empirical study. We conclude that a long-term effect of a reduction in the density of planktivorous fish can be expected only when the external phosphorus loading is reduced to below 0.5–2.0 g m⁻² y⁻¹. This loading is equivalent to an in-lake summer concentration below 80–150μg P l⁻¹. Furthermore, fish manipulation as a restoration tool seems most efficient in shallow lakes.     

Temporal and geographical variation in lake trophic status in the English Lake District: evidence from (sub)fossil diatoms and aquatic macrophytes

1. A sediment core (representing 250–300 years) was taken from each of three lakes of conservation interest and contrasting trophic status in the English Lake District: Wastwater, Bassenthwaite Lake and Esthwaite Water. Lithostratigraphic analyses, radiometric dating and analysis of fossil diatoms were carried out.
2. Transfer functions, based on the diatoms, were used to reconstruct total phosphorus (TP) and, thus, eutrophication at the study lakes. In Wastwater, changes in lake pH were also reconstructed.
3. The lakes were also classified according to their present macrophyte flora, the latter being compared with previous records.
4. The fossil diatoms of Wastwater were continuously dominated by taxa typical of oligotrophic, circumneutral waters, indicating that the lake has not been enriched or acidified in the last 250 years. The aquatic macrophyte flora has probably remained unchanged since before the Industrial Revolution.
5. The diatom assemblages of both Bassenthwaite Lake and Esthwaite Water began to change in the mid-1800s. Further change occurred from the 1960s, at the onset of a recent period of eutrophication. These two lakes have experienced continued nutrient enrichment throughout the 1970s, 80s and 90s, largely associated with increasing phosphorus inputs from sewage effluent. There is no evidence of any recovery in response to recent reductions in external nutrient loads.
6. Only in Esthwaite Water has the change in aquatic macrophytes been pronounced.
7. Palaeolimnological reconstruction is useful in determining background conditions and natural variation in lake ecosystems.     

Modelling nutrient cycles in relation to food web structure in a biomanipulated shallow lake

The modelPCLAKE describes the phosphorus and nitrogen cycles within a shallow lake ecosystem, including the sediment and a simplified biological food web. All components are modelled in a generalized way rather than a very detailed one. This model has been applied to Lake Zwemlust, a small biomanipulated lake in The Netherlands. Formerly, this highly eutrophic lake was dominated by cyanobacteria and devoid of macrophytes. Biomanipulation was carried out in 1987 by pumping-out of the water, removal of all fish, and refilling of the lake with seepage water. The lake was restocked with some rudd, pike, zooplankton and seedlings of macrophytes, and then monitored up to 1992. Macrophytes developed rather quickly and reached their maximum biomass during the six-years period in 1989. Despite the continuously high nutrient (N and P) loading, algal biomass remained low due to nitrogen limitation, caused by competition with the macrophytes. From 1990 onwards, the macrophytes declined again and a species shift occurred, following an increase of herbivorous birds on the lake and the development of herbivorous fishes.Model simulations grossly reproduced the observed developments in Lake Zwemlust before and after the biomanipulation measures. The existence of multiple steady states at the same trophic state and the possible shift between them could be simulated well. This study also demonstrates the interrelation between system structure and the distribution and cycling of nutrients. It is concluded, that within general boundary conditions set by the trophic state of the system, the food web structure determines the actual nutrient flows and the occurrence of nutrient limitations of the primary producers. It is shown that both aspects can be integrated in one mathematical model. The long-term stability of the macrophyte dominance in the lake is discussed.     

Rotifer community structure in three shallow lakes: seasonal fluctuations and explanatory factors

The present work aimed at studying the rotifer communities of three shallow eutrophic lakes in Portugal (lakes Mira, Vela and Linhos). At the time of the study, Mira and Vela faced large inputs of allochthonous nutrients, while Linhos was facing terrestrialisation, with cycles of dominance-senescence of macrophytes. The three lakes differed in terms of their abiotic features, with Linhos presenting very high nutrient levels and low pH, while Vela and Mira shared most of the characteristics. The rotifer communities of these two lakes were poorly diversified but highly abundant (max. > 2000 ind l⁻¹), with a clear dominance of eurytopic euplanktonic species (mainly Keratella cochlearis). On the other hand, Linhos presented lower abundances (<1000 ind l⁻¹) but higher species richness, mainly due to macrophyte-associated taxa, such as the littoral genera Lepadella, Testudinella and Squatinella. In all lakes, summertime represented a peak in terms of abundance and diversity. Canonical correspondence analysis (CCA) identified two main environmental gradients that shape up the rotifer assemblages: a temporal gradient, mainly related to temperature, and a eutrophy gradient, associated with nitrogenous nutrients. The latter gradient is clearly dependent on between-lake variation, due to the high nutrient levels observed in lake Linhos. Variance partitioning using CCA revealed that the largest portion (27.5%) of the total variation explained (52.1%) was attributed to the interaction between lake and environmental variables.     

Back to the future: using palaeolimnology to infer long‐term changes in shallow lake food webs

1. Shallow lakes are often cited as classic examples of systems that exhibit trophic cascades but, whilst they provide good model systems with which to test general ecological theory and to assess long‐term community change, their food web linkages have rarely been resolved, so changes associated with the structure and dynamics of the ecological network as a whole are still poorly understood. 2. We sought to redress this, and to demonstrate the potential benefits of integrating palaeolimnological and contemporary data, by constructing highly resolved food webs and stable isotope derived measures of trophic interactions and niche space, for the extant communities of two shallow U.K. lakes from different positions along a gradient of eutrophication. The contemporary surface sediment cladoceran and submerged macrophyte assemblages in the less enriched site, Selbrigg Pond, matched the palaeolimnological assemblages of the more enriched site, Felbrigg Hall Lake, in its more pristine state during the 1920s. Thus, Selbrigg was a temporal analogue for Felbrigg, from which the consequences of long‐term eutrophication on food web structure could be inferred. These data represent the first steps towards reconstructing not only past assemblages (i.e. nodes within a food web), but also past interactions (i.e. links within a food web): a significant departure from much of the previous research in palaeolimnology. 3. The more eutrophic food web had far fewer nodes and links, and thus a less reticulate network, than was the case for the more pristine system. In isotopic terms, there was vertical compression in δ¹⁵N range (NR) and subsequent increased overlap in isotopic niche space, indicating increased trophic redundancy within Felbrigg. This structural change, which was associated with a greater channelling of energy through a smaller number of nodes as alternative feeding pathways disappear, could lead to reduced dynamic stability, pushing the network towards further simplification. These changes reflected a general shift from a benthic‐dominated towards a more pelagic system, as the plant‐associated subweb eroded. 4. Although these data are among the first of their kind, the palaeo‐analogue approach used here demonstrates the huge potential for applying food web theory to understand how and why these ecological networks change during eutrophication. Furthermore, because of the rich biological record preserved in their sediments, shallow lakes represent potentially important models for examining long‐term intergenerational dynamics, thereby providing a means by which models and data can be integrated on meaningful timescales – a goal that has long proved elusive in food web ecology.