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Abundances of different functional groups of waterfowl in L. Krankesjön during May–October from 1985 to 2007 (bars), and regional abundance index for summer and winter countings (line). Species include herbivores: Mute Swan (Cygnus olor), Coot (Fulica atra), and Eurasian Wigeon (Anas penelope); invertebrate feeders: Goldeneye (Buchepala clangula) and tufted duck (Aythia fuligula); fish feeders: Great crested grebe (Podiceps cristatus) and Common merganser (Mergus merganser); and omnivorous species: (Common pochard (Aythia ferina)), as well as waterfowl, mainly feeding outside the lake including Mallard (Anas platyrhynchos) and Teal (Anas crecca)  

Abundances of different functional groups of waterfowl in L. Krankesjön during May–October from 1985 to 2007 (bars), and regional abundance index for summer and winter countings (line). Species include herbivores: Mute Swan (Cygnus olor), Coot (Fulica atra), and Eurasian Wigeon (Anas penelope); invertebrate feeders: Goldeneye (Buchepala clangula) and tufted duck (Aythia fuligula); fish feeders: Great crested grebe (Podiceps cristatus) and Common merganser (Mergus merganser); and omnivorous species: (Common pochard (Aythia ferina)), as well as waterfowl, mainly feeding outside the lake including Mallard (Anas platyrhynchos) and Teal (Anas crecca)  

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The importance of lake ecosystems for waterfowl remains a topic of debate. In order to assess how temporal variations in lake features, specifically shifts between alternative stable states, may interact with the waterfowl fauna, we performed a long-term (22years) study of the shallow Lake Krankesjön, southern Sweden. Lower total numbers of waterfo...

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... Long-term studies of the submerged macrophytes at Lake Krankesjön, Sweden also strongly suggest that waterbird herbivory on Chara spp. has no long-term effects on species composition, persistence or biomass of these macrophytes at this site (Hansson et al., 2010). Seventeen large-scale waterbird exclosure plots monitored over 2 years at Lake Botshol, Netherlands showed no significant increase in Charales biomass in the absence of grazing compared with the controls during transition from turbid to clear water and high Chara biomass, demonstrating that light limitation was the main factor controlling the collapse and return of Charales there (Rip, Rawee & de Jong, 2006). ...
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Stoneworts (Charales) are green algae that represent an important food resource for many waterbird species in Europe and elsewhere. Browsing avian herbivores (e.g. swan, goose, duck and coot species) consume Charales plant vegetative parts, by head‐dipping, up‐ending or diving. A lower fibre content and longer growing season may make Charales as attractive to such herbivores as sympatric submerged higher plant species in some circumstances. Charales respond to environmental stress (e.g. drought) by producing abundant diaspores, in the form of oospores (sexual) and bulbils (asexual), both rich in starch, generating abundant food for waterbirds at critical stages in their annual migratory cycles. Waterbirds feed on these by diving (e.g. common pochard Aythya ferina and red‐crested pochard Netta rufina ) or by filtering from the water column (e.g. dabbling duck species), ensuring dispersal of sexually produced and vegetative diaspores locally (because of predator swamping) and remotely (through endo‐ and ectozoochorous dispersal by long‐distance migratory waterbirds). Greater invertebrate density and diversity associated with Charales canopies enhances their attractiveness over other submerged macrophyte beds to diving predators [e.g. tufted duck Aythya fuligula , common pochard and Eurasian coot Fulica atra (hereafter coot)]. Fish fry preying on these invertebrates use such vegetation as predator cover, in turn providing prey for avian piscivores such as grebes and cormorants. Abundant Charales contribute to maintaining a transparent water column due to canopy density, nutrient effects, dampening of sedimentation/remobilisation of suspended matter and nutrients and allelopathic effects on other plants (especially phytoplankton). Shallow, relatively eutrophic waters can flip between clear‐water high‐biodiversity (where Charales thrive) and turbid species‐poor depauperate stable states (lacking Charales). Shifts between turbid conditions and rich submerged Charales beds have profound elevating effects on aquatic diversity, to which waterbirds show rapid aggregative responses, making them ideal indicator species of ecological change; in the case of Charales specialists (such as red‐crested and common pochard), indicators of the presence and abundance of these plants. Large‐bodied colonial nesting birds (e.g. cormorants, gulls, heron and egrets) aggregating along lake shores contribute high N and P loadings to water bodies sensitive to such external and internal inputs and can cause local eutrophication and potential loss of Charales. Despite variation from complete seasonal removal of Charales biomass to undetectable grazing effects by herbivorous birds, evidence suggests little effect of avian grazing on biomass accumulation or the stability of community composition (under otherwise stable conditions), but we urge more research on this under‐researched topic. We also lack investigations of the relative foraging profitability of different Charales organs to waterbirds and the degree of viability of gyrogonites (fertilised and calcified oospores), vegetative bulbils and plant fragments after passage through the guts of waterbirds. We especially need to understand better how much the carbonate armour of these organs affects their viability/dispersal via waterbirds and urge more research on these neglected plants and their relationships and interactions with other organisms in the aquatic biota.
... Shallow lakes are highly productive ecosystems that support a major aquatic and terrestrial biodiversity, even despite the small dimensions of some lakes, but they maintain a fragile balance that can be altered by an increase in nutrients leading to a regime shift, with a high loss of biodiversity and deterioration of ecosystem services (Declerck et al. 2005). Shallow wetlands are especially important in bird conservation because of their crucial role in supporting waterbirds populations and biodiversity, or as favourable migration or breeding habitat for many bird species (Kleijn et al. 2014;Zhang et al. 2015;Gaget et al. 2020), although the influence of birds on the trophic status of shallow wetlands is still widely unknown (Dejoux 1983;Gere and Andrikovics 1992;Suter 1994;Hansson et al. 2010). ...
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... Shallow lakes are highly productive ecosystems that support a major aquatic and terrestrial biodiversity, even despite the small dimensions of some lakes, but they maintain a fragile balance that can be altered by an increase in nutrients leading to a regime shift, with a high loss of biodiversity and deterioration of ecosystem services (Declerck et al. 2005). Shallow wetlands are especially important in bird conservation because of their crucial role in supporting waterbirds populations and biodiversity, or as favourable migration or breeding habitat for many bird species (Kleijn et al. 2014;Zhang et al. 2015;Gaget et al. 2020), although the influence of birds on the trophic status of shallow wetlands is still widely unknown (Dejoux 1983;Gere and Andrikovics 1992;Suter 1994;Hansson et al. 2010). ...
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... In Lake Rautajärvi, the reduction in the number of pairs has been more regular, and in the study period, we calculated a 46.5% We hypothesized that water browni cation is the main reason explaining the decrease in the density of waterbirds. It seems that browni cation has an in uence on water quality and, consequently, on waterbirds (Hansson et al. 2010, Blanchet et al. 2022). COD has traditionally been considered as a parameter for the estimation of organic matter in water (Sepp et al. 2018). ...
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Brownification of freshwater is a global issue with many implications. The high accumulation of organic matter causing the darkening of freshwater is explained by numerous factors including climate and land use. One example of this alteration can be found in the population of waterbirds (fish eaters, diving and dabbling ducks and other species) in lakes Kukkia and Rautajärvi in Finland, where the number of waterbirds has decreased by 40% since the mid-1980s. We run Generalized Linear Mixed Models to determine how climatic, water chemistry and land use factors are interacting with one another and how those interactions impact waterbird density. Our results showed the significance of brownification and climatic factors, as COD (chemical oxygen demand) and turbidity of the water body and wind speed were the most relevant in explaining the decline of waterbirds.
... Predatory impact of piscivorous birds on the trophic relationships in temperate standing waters seems to be more diffuse or negligible (Paszkowski and Tonn, 2000). Too much waterbirds can shift SSWs in a turbid state, (Hansson et al., 2010). They can also be strong contributors to the dissemination of exotic invertebrate and plant species (Brochet et al., 2009;Frisch et al., 2007). ...
... They can also be strong contributors to the dissemination of exotic invertebrate and plant species (Brochet et al., 2009;Frisch et al., 2007). Because eutrophication also favours high abundance of omnivorous waterbirds (Hansson et al., 2010), it may therefore be difficult to disentangle waterbird from eutrophication impacts. Invertivorous or opportunistic exotic fish species can strongly modify invertebrate communities (e.g. ...
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Small shallow lakes and ponds (SSWs) have high conservation value and support numerous ecosystem services. However, these small ecosystems face many threats, including eutrophication and internal biotic pressures like Cyprinidae or exotic bioturbators crayfish, which tend to shift biodiverse SSWs to a turbid state dominated by phytoplankton unfitted to numerous usages, and compromise the effectiveness of their restoration. The ecological quality of SSWs and the efficiency of their management still remain poorly evaluated because of the lack of adapted tools. To fill this gap, we propose a new multimetric index (BECOME) and a diagnostic tool (BECOMEd) both based on macrophyte and invertebrate communities. BECOME exhibits a high sensitivity to the impact of surrounding crops, urbanization and fertilized meadows, morphological alterations, and various internal biotic sources of alterations, in a wide variety of geological and climatic contexts. BECOME is especially innovative, because taking into account internal biotic sources of pressures neglected by most of the existing biological WFD-compliant indices for European lakes and ponds, despite their importance in SSW threats. BECOMEd allows to identify the major alteration sources and estimates their relative contribution to SSW degradation, helping managers to prioritize actions or to evaluate their effectiveness. The statistical design applied for the index construction has been fitted to develop biological index for ecosystems influenced by a wide amplitude of numerous environmental conditions. This design could be easily applied elsewhere in the world, for SSWs but also for other types of ecosystems.
... Possibly, in invaded ponds macroinvertebrates shift towards the green path due either to a lower use of this resource by vertebrates or a displacement of invertebrates from the brown path by vertebrates because of competition, direct consumption, predation avoidance behavior or direct interferences. Indeed, given that most algae on these systems grow attached to macrophytes, a displacement of macroinvertebrates from the bottom to macrophyte patches-seeking refuge from predation or because bioturbation-may promote a diet including more algal resources (Hansson et al. 2010). ...
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Alien species introductions produce strong impacts on invaded communities, altering their structure, diversity and functioning. These impacts are interrelated with changes in food web architecture. However, the reorganization or robustness of food webs in the face of invasion is a phenomenon poorly considered in ecology and conservation practices. In this article, we analyze the effects of the invasion of the American bullfrog, Lithobates catesbeianus, on the structure and function of invaded food webs. Specifically, we analyzed the integration of energetic channels by top predators, the relative use of alternative energetic paths by different functional groups and its dependence on body size among invaded and uninvaded communities in Uruguay, South America. The integration of energetic paths at high trophic positions by large body sized consumers was a pervasive feature of food webs among all studied ponds, in spite of turnover in top predator identity and large changes in community composition. Bullfrog post-metamorphs presented high trophic positions, integrating the primary producers and detritus paths, acting as apex predators in invaded food webs. The bullfrog tadpoles presented intermediate positions and were associated with the detritivorous pathway. On the other hand, the relative importance of the primary producers and detritus as carbon sources assimilated into the biomass of fish and invertebrates was altered in invaded systems. The robustness in the integration pattern of energy channels is congruent with its proposed central role in the stability of food webs. These results advance the understanding of the effect of invasions on key structural features of food webs, notably underrepresented in the invasion literature.
... By doing so, it is expected that it will facilitate various primary producers that stimulate the development of the food web in the lake to benefit the higher trophic levels, including fish and water birds (Hansson et al., 2010;van Leeuwen et al., 2021). Our experiments show that, provided that (a) this new archipelago generates sufficient shelter and (b) that herbivory pressure by waterfowl is not too high (Bakker, Sarneel, et al., 2013;Bakker et al., 2016), the benthic aquatic ecosystem of the Marker Wadden that is protected by the archipelago may become dominated by submerged macrophytes over time. ...
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Wind‐induced turbulence can strongly impact ecological processes in shallow lake ecosystems. The creation of shelter against wind can be expected to affect both primary producers and herbivores in aquatic food webs. Shelter may benefit particular primary producers more than others by changing relative resource availabilities for different primary producers. Herbivore community compositions may be affected either directly or indirectly as a consequence of changes in their food quantity and quality that, in turn, may affect the transfer efficiency between primary producers and herbivores. A reduction in trophic transfer efficiency resulting from wind‐induced turbulence potentially can lead to declines of higher trophic levels, but is generally understudied. Here, we focus on the impact of wind on aquatic primary producers and trophic transfer efficiency. We hypothesised that reducing wind‐induced turbulence will stimulate higher trophic production in shallow lakes. However, the multitude of impacts of wind‐induced turbulence on aquatic food webs make it challenging to predict the direction of change when creating sheltered conditions. We tested our hypothesis in the shallow waters of a newly constructed archipelago named Marker Wadden in lake Markermeer in the Netherlands. Lake Markermeer has experienced declining numbers of birds and fish. These declines have been related to wind‐induced sediment resuspension that potentially limits primary production and trophic transfer efficiency. Marker Wadden is a large‐scale restoration project that aims to add sheltered and heterogeneous habitat to the otherwise mostly homogeneous lake, thus targeting the potential problems associated with wind‐induced turbulence. We executed a 2‐month manipulative field mesocosm experiment in the shallow waters of Marker Wadden to study the effect of reduced wind‐induced turbulence (i.e., shelter) on aquatic food webs. Specifically, we studied the effects on primary producers, trophic transfer efficiency between phytoplankton and zooplankton (using zooplankton biomass divided by phytoplankton Chl a as a proxy), and benthic fauna. The experiment consisted of three treatments: no shelter , shelter without macrophytes and shelter with submerged macrophytes ( Myriophyllum spicatum ) present at the start of the experiment . Our results clearly showed that under unsheltered conditions phytoplankton was the dominant primary producer, whereas in sheltered conditions submerged macrophytes became dominant. Interestingly, submerged macrophytes appeared rapidly in the sheltered treatment where first no macrophytes were visibly present; hence, at the end of the experiment, there was little difference among the sheltered treatments with and without initial presence of submerged macrophytes. Despite that phytoplankton concentrations were 23‐fold higher under the unsheltered conditions, this did not result in higher zooplankton biomass. This can be explained by a five‐fold greater trophic transfer efficiency between phytoplankton and zooplankton under the sheltered conditions. Furthermore, under the sheltered conditions the Gastropoda density reached 746 individuals m ⁻² , whereas no Gastropoda were found under the no shelter treatment. These findings indicate that for shallow lakes that are negatively affected by wind‐induced turbulence, measures aimed at ameliorating this stressor can be effective in facilitating submerged macrophyte recovery, increasing Gastropoda densities and restoring trophic transfer efficiency between phytoplankton and zooplankton. Ultimately, this may support higher trophic levels such as fish and water birds by increasing their food availability in shallow lake ecosystems.
... In the boreal environment, the abundance of waterbirds feeding on fishes, plants and invertebrates showed a positive relation with clear water and macrophyte percentage cover (Hansson et al., 2010). The impact of browning on waterbirds has not been demonstrated yet, but water browning causes macrophyte decline (Reitsema et al., 2018). ...
Article
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Water browning or brownification refers to increasing water color, often related to increasing dissolved organic matter (DOM) and carbon (DOC) content in freshwaters. Browning has been recognized as a significant physicochemical phenomenon altering boreal lakes, but our understanding of its ecological consequences in different freshwater habitats and regions is limited. Here, we review the consequences of browning on different freshwater habitats, food webs and aquatic-terrestrial habitat coupling. We examine global trends of browning and DOM/DOC, and the use of remote sensing as a tool to investigate browning from local to global scales. Studies have focused on lakes and rivers while seldom addressing effects at the catchment scale. Other freshwater habitats such as small and temporary waterbodies have been overlooked, making the study of the entire network of the catchment incomplete. While past research investigated the response of primary producers, aquatic invertebrates and fishes, the effects of browning on macrophytes, invasive species, and food webs have been understudied. Research has focused on freshwater habitats without considering the fluxes between aquatic and terrestrial habitats. We highlight the importance of understanding how the changes in one habitat may cascade to another. Browning is a broader phenomenon than the heretofore concentration on the boreal region. Overall, we propose that future studies improve the ecological understanding of browning through the following research actions: 1) increasing our knowledge of ecological processes of browning in other wetland types than lakes and rivers, 2) assessing the impact of browning on aquatic food webs at multiple scales, 3) examining the effects of browning on aquatic-terrestrial habitat coupling, 4) expanding our knowledge of browning from the local to global scale, and 5) using remote sensing to examine browning and its ecological consequences.
... According to this study, fresh to brackish SAV species rarely considered in coastal management and restoration decision-making (CPRA, 2017;Wilson et al., 2002) provide an extensive area of habitat (food and cover) and occupy a meaningful role in the estuarine ecosystems in the NGOM region. Including these habitats and SAV species in assessment, ecosystem management, and restoration could benefit coastal and estuarine managers as these habitats have been shown to dramatically impact wildlife populations and ecosystem services (Brasher et al., 2012;Castellanos & Rozas, 2001;Hansson et al., 2010;Hitch et al., 2011;Kanouse et al., 2006;La Peyre & Gordon, 2012). Specifically, incorporation of these occurrence and distribution estimates of this foundational species are necessary to consider in predictions of the possible future coastal landscape as water bodies with SAV present provide different ecological benefits (i.e., sediment stabilization, nutrient cycling, impacts to water movement) than those without, as well as provide more accurate estimates of fish and wildlife populations (food and habitat provisioning). ...
Article
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... 7−10 Aquatic macrophytes are recognized for their ecosystem services in shallow freshwater ecosystems. 11,12 When they die and decompose, vast concentrations of nutrients are released from plant litter, resulting in seasonal deterioration of the water quality. 13 In aquatic ecosystems, macrophyte-derived detritus decomposition is influenced not only by internal factors, such as initial litter quality, but also by external environmental factors such as water temperature, 14−17 composition of the decomposer community, 14,18 and nutrient availability in the water column, 19 as well as in the sediment. ...
... The enclosures were randomly distributed on a flat area and filled with 400 L of unfiltered lake water collected from the eutrophic Lake Krankesjon (N55°42′, E13°27′), a shallow lake located in Southern Sweden. For a more thorough lake description, consider Hansson et al. 11 Our study consisted of three temperature treatments (each replicated eight times): controls, mimicking the current climate state in a temperate shallow lake in southern Sweden (hereafter named U treatment: unheated treatment); a treatment where the temperature followed the ambient daily and seasonal variations, but at a 4°C higher level (hereafter named W treatment: warmed); and finally, a treatment with a preprogrammed fluctuating temperature, ranging from 0 to 8°C above ambient conditions (hereafter named H treatment: heat waves), mimicking the predicted future climate scenario of more frequent and intense temperature variations. 4 The frequency and amplitude of the heat waves were based on model predictions from IPCC (2013) and the Swedish Meteorological and Hydrological Institute (SMHI) for a climate scenario about 75 years into the future. ...
Article
In addition to a rise in global air and water mean temperatures, extreme climate events such as heat waves are increasing in frequency, intensity, and duration in many regions of the globe. Developing a mechanistic understanding of the impacts of heat waves on key ecosystem processes and how they differ from just an increase in mean temperatures is therefore of utmost importance for adaptive management against effects of global change. However, little is known about the impact of extreme events on freshwater ecosystem processes, particularly the decomposition of macrophyte detritus. We performed a mesocosm experiment to evaluate the impact of warming and heat waves on macrophyte detrital decomposition, applied as a fixed increment (+4 °C) above ambient and a fluctuating treatment with similar energy input, ranging from 0 to 6 °C above ambient (i.e., simulating heat waves). We showed that both warming and heat waves significantly accelerate dry mass loss of the detritus and carbon (C) release but found no significant differences between the two heated treatments on the effects on detritus dry mass loss and C release amount. This suggests that moderate warming indirectly enhanced macrophyte detritus dry mass loss and C release mainly by the amount of energy input rather than by the way in which warming was provided (i.e., by a fixed increment or in heat waves). However, we found significantly different amounts of nitrogen (N) and phosphorus (P) released between the two warming treatments, and there was an asymmetric response of N and P release patterns to the two warming treatments, possibly due to species-specific responses of decomposers to short-term temperature fluctuations and litter quality. Our results conclude that future climate scenarios can significantly accelerate organic matter decomposition and C, N, and P release from decaying macrophytes, and more importantly, there are asymmetric alterations in macrophyte-derived detrital N and P release dynamic. Therefore, future climate change scenarios could lead to alterations in N/P ratios in the water column via macrophyte decomposition processes and ultimately affect the structure and function of aquatic ecosystems, especially in the plankton community.