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Shallow lakes are a key component of the global carbon cycle. It is, therefore, important to know how shallow lake ecosystems will respond to the current climate change. Global warming affects not only average temperatures, but also the frequency of heat waves (HW). The impact of extreme events on ecosystems processes, particularly greenhouse gas ( GHG) emissions, is uncertain., Using the world's longest-running shallow lake experiment, we studied the effects of a simulated summer HW on the fluxes of carbon dioxide ( CO2), methane ( CH4) and nitrous oxide (N2O). The experimental mesocosms had been exposed to different temperature treatments and nutrient loading for 11 years prior to the artificial HW., In general, there was an increase in total GHG emissions during the 1-month artificial HW, with a significant increase in CO2, CH4 and N2O being observed in the shallow lake mesocosms. No significant effect of the HW on CO2 emissions could be traced, though, in the mesocosms with high nutrient levels. Furthermore, the data suggested that in addition to the direct effect of increased temperature on metabolic processes during the HW, biotic interactions exerted a significant control of GHG emissions. For example, at low nutrient levels, increased CO2 emissions were associated with low macrophyte abundance, whereas at high nutrient levels, decreased phytoplankton abundance was linked to increased emissions of CO2 and CH4., In contrast to the observable heat-wave effect, no clear general effect of the long-term temperature treatments could be discerned over the summer, likely because the potential effects of the moderate temperature increase, applied as a press disturbance, were overridden by biotic interactions. This study demonstrates that the role of biotic interactions needs to be considered within the context of global warming on ecosystem processes. [ABSTRACT FROM AUTHOR]

1. Seasonal relationships between macrophyte and phytoplankton populations may alter considerably as lakes undergo eutrophication. Understanding of these changes may be key to the interpretation of ecological processes operating over longer (decadal-centennial) timescales. 2. We explore the seasonal dynamics of macrophytes (measured twice in June and August) and phytoplankton (measured monthly May–September) populations in 39 shallow lakes (29 in the U.K. and 10 in Denmark) covering broad gradients for nutrients and plant abundance. 3. Three site groups were identified based on macrophyte seasonality; 16 lakes where macrophyte abundance was perennially low and the water generally turbid (‘turbid lakes’); 7 where macrophyte abundance was high in June but low in August (‘crashing’ lakes); and 12 where macrophyte abundance was high in both June and August (‘stable’ lakes). The seasonal behaviour of the crashing and turbid lakes was extremely similar with a consistent increase in nutrient concentrations and chlorophyll- a over May–September. By contrast in the stable lakes, seasonal changes were dampened with chlorophyll- a consistently low (<10–15 μg L−1) over the entire summer. The crashing lakes were dominated by one or a combination of Potamogeton pusillus, Potamogeton pectinatus and Zannichellia palustris, whereas Ceratophyllum demersum and Chara spp. were more abundant in the stable lakes. 4. A long-term loss of macrophyte species diversity has occurred in many shallow lakes affected by eutrophication. One common pathway is from a species-rich plant community with charophytes to a species-poor community dominated by P. pusillus, P. pectinatus and Z. palustris. Such compositional changes may often be accompanied by a substantial reduction in the seasonal duration of plant dominance and a greater tendency for incursions by phytoplankton. We hypothesise a slow-enacting (10–100 s years) feedback loop in nutrient-enriched shallow lakes whereby increases in algal abundance are associated with losses of macrophyte species and hence different plant seasonal strategies. In turn such changes may favour increased phytoplankton production thus placing further pressure on remaining macrophytes. This study blurs the distinction between so-called turbid phytoplankton-dominated and clear plant-dominated shallow lakes and suggests that plant loss from them may be a gradual process. [ABSTRACT FROM AUTHOR]

1. Quantitative palaeolimnology has traditionally sought to quantify species-environment relationships to use alterations in biological assemblages to reflect past environmental change. Transfer functions have used regression techniques, such as weighted averaging, to define taxon optima and tolerance for a single chemical or biological variable. 2. Cladoceran assemblages and their sub-fossil remains in shallow lakes are shaped by a combination of interacting factors. Partial constrained ordination of sub-fossil cladoceran assemblages from 39 shallow lakes (29 in Norfolk, U.K. and 10 in Denmark) indicated that both zooplanktivorous fish (ZF) density and submerged macrophyte abundance significantly influenced community composition. These dual structuring forces precluded the use of a transfer function as one of the key assumptions of this approach was not met, namely that environmental variables apart from the variable being modelled have negligible influence on species distribution or that there is a linear relationship between the two. Separate transfer functions for ZF and macrophyte abundance were developed but had poor performance diagnostics with low bootstrapped r2, high root mean square error of prediction (RMSEP) and large bias. 3. To obviate the problem of multiple structuring forces a multivariate regression tree (MRT) was employed, which allows for more than one explanatory variable within a model. The MRT analysis defined six groups with discrete ranges of ZF and macrophyte densities. The technique identified critical values or ‘break points’ in ZF and macrophyte abundances which result in significant alterations in the sub-fossil cladoceran assemblage. In addition, the MRT groups had different summer mean values for chlorophyll-a, Secchi depth, total phosphorus and nitrate-nitrogen. 4. The predictive abilities of the model were assessed by comparing the observed versus predicted MRT group membership. In general group membership was reliably predicted, suggesting sub-fossil cladoceran assemblages reliably reflect ZF and macrophyte density in shallow lakes. For a relatively small number of sites there were differences between the observed and predicted MRT group membership. These failures of prediction may result, at least in part, from the disparity of the time period represented by the environmental data and the surface sediment cladoceran assemblage. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

1. Eutrophication has a profound effect on the biological structure and function of shallow lakes, altering the composition and abundance of submerged macrophyte and fish assemblages. Relatively little is known, however, about decadal to centennial-scale change in these important aspects of shallow lake ecology. 2. Established palaeolimnological inference models are limited to reconstructing a single variable. As macrophyte and zooplanktivorous fish abundance exert dual and interacting controls on cladoceran assemblages a single variable inference model may contain significant error. To obviate this problem, we applied a new cladoceran-based multivariate regression tree (MRT) model to cladoceran subfossil assemblages from dated cores from a small shallow lake (Felbrigg Lake, U.K.) to assess long-term change in fish and submerged macrophyte abundance. Plant macrofossil, chironomid and mollusc subfossil assemblages were also analysed to track changes in biological structure and function and to evaluate the inferences of the MRT model. 3. Over the 200+ year period covered by the sediment cores, there was good agreement in the timing and nature of ecological change reflected by the plant macrofossil, mollusc, chironomid and cladoceran data. The sediment sequence was divided into three dated zones: c. 1797–1890, c. 1890–1954 and c. 1954–present. Prior to 1890 plant-associated mollusc, cladoceran and chironomid assemblages indicated a species-rich macrophyte community; a scenario confirmed by the plant macrofossil data. From c. 1890 to 1954 macrophyte-associated species of all three invertebrate groups remained abundant but the proportion of pelagic cladocerans rose. Post-1954 mollusc and chironomid assemblages changed to sediment associated detrital feeders and the proportion of pelagic cladoceran taxa increased further. 4. The cladoceran-based MRT model indicated a long period of stability, c. 1790–1927, characterised by abundant submerged macrophytes and zooplanktivorous fish. From c. 1927 to 1980, the MRT model inferred a decline in zooplanktivorous fish density (ZF) but relative stability in August macrophyte abundance. From 1980 to 2000, an increase in zooplanktivorous fish was inferred tallying well with available data on the fish population (since the 1970s), which indicated extirpation of perch in the 1970s and a subsequent increase in the rudd population. The model inferred little change in August macrophyte abundance until post- c. 1980 at which point it indicated a decline. The surface sediment assemblage was placed in MRT group A, where submerged plants are absent or very rare in late summer in good agreement with current conditions at the site. 5. The MRT model, applied here for the first time, appears to have successfully tracked changes in macrophyte abundance and ZF over the last 200 years at Felbrigg Lake. The inferences agreed with historical observations on the fish community and the supporting palaeolimnological data. Given that multiple structuring forces shape most biological communities, the application of a model capable of allowing for this represents a significant advance in palaeolimnology. [ABSTRACT FROM AUTHOR]

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 undertwinspan 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. [ABSTRACT FROM AUTHOR]

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. [ABSTRACT FROM AUTHOR]

Shallow lakes are globally the most numerous water bodies and are sensitive to external perturbations, including eutrophication and climate change, which threaten their functioning. Extreme events, such as heat waves (HWs), are expected to become more frequent with global warming. To elucidate the effects of nutrients, warming, and HWs on zooplankton community structure, we conducted an experiment in 24 flow-through mesocosms (1.9 m in diameter, 1.0 m deep) imitating shallow lakes. The mesocosms have two nutrient levels (high (HN) and low (LN)) crossed with three temperature scenarios based on the Intergovernmental Panel on Climate Change (IPCC) projections of likely warming scenarios (unheated, A2, and A2 + 50%). The mesocosms had been running continuously with these treatments for 11 years prior to the HW simulation, which consisted of an additional 5 °C increase in temperature applied from 1 July to 1 August 2014. The results showed that nutrient effects on the zooplankton community composition and abundance were greater than temperature effects for the period before, during, and after the HW. Before the HW, taxon richness was higher, and functional group diversity and evenness were lower in HN than in LN. We also found a lower biomass of large Cladocera and a lower zooplankton: phytoplankton ratio, indicating higher fish predation in HN than in LN. Concerning the temperature treatment, we found some indication of higher fish predation with warming in LN, but no clear effects in HN. There was a positive nutrient and warming interaction for the biomass of total zooplankton, large and small Copepoda, and the zooplankton: phytoplankton ratio during the HW, which was attributed to recorded HW-induced fish kill. The pattern after the HW largely followed the HW response. Our results suggest a strong nutrient effect on zooplankton, while the effect of temperature treatment and the 5 °C HW was comparatively modest, and the changes likely largely reflected changes in predation. [ABSTRACT FROM AUTHOR]

Shallow lakes produce and emit substantial amounts of methane (CH 4). Part of the CH 4 produced in lakes is consumed by methane-oxidizing bacteria (MOB) present in the sediment and water column, thus reducing the overall CH 4 emissions. However, the role of aquatic plants as habitat for CH 4 oxidation by MOB is poorly understood. In this study, we compared CH 4 oxidation rates and MOB abundance associated with different types of aquatic plants (periphyton, filamentous algae, and both above-ground macrophytes and their rhizosphere). The plants were collected from shallow lake mesocosms exposed to experimental nutrient enrichment and warming treatments for 17 years prior to this study. Incubations of all sampled plants showed CH 4 oxidation, with above-ground macrophyte tissue and filamentous algae having the highest rates of up to 0.25 µmol CH 4 h−1 g−1 dw. Oxidation rates associated with macrophytes were species dependent, with consumption rates on rhizospheres of Potamogeton crispus higher than those on Elodea canadensis. The increase in nutrients and dissolved CH 4 in the water tended to increase MOB abundance and activity for all plant types, while no effect of long term warming was detectable. Our results showed that MOB associated with periphyton, filamentous algae and macrophytes oxidize CH 4 in shallow lakes at different rates across species or plant types. We also found that high macrophyte biomass is associated with reduced CH 4 concentration in the water. This study shows that CH 4 oxidation occurs on many plant surfaces but that oxidation rates alone cannot explain the reduced CH 4 emissions at higher plant biomass. [ABSTRACT FROM AUTHOR]