Fish communities face increasing anthropogenic pressures in freshwater and marine ecosystems that modify their biodiversity and threaten the services they supply to human populations. To address ...these issues, studies have been increasingly focusing on functions of fish that are linked to their main ecological roles in aquatic ecosystems. Fish are indeed known to control other organisms through predation, mediate nutrient fluxes, and can act as ecosystem engineers. Here for each of the key functions played by fish, we present the functional traits that have already been used to assess them. We include traits measurable from observations on living individuals, morphological features measured on preserved organisms or traits categorized using information from the literature, and we discuss their respective advantages and limitations. We then list future research directions to foster a more complete functional approach for fish ecology that needs to incorporate functional traits describing, food provisioning and cultural services while accounting more frequently for intraspecific variability. Finally, we highlight ecological and evolutionary questions that could be addressed using meta-analyses of large trait databases, and how a trait-based framework could provide valuable insights on the mechanistic links between global changes, functional diversity of fish assemblages, and ecosystem services.
ABSTRACT
The role of animals in modulating nutrient cycling hereafter, consumer‐driven nutrient dynamics (CND) has been accepted as an important influence on both community structure and ecosystem ...function in aquatic systems. Yet there is great variability in the influence of CND across species and ecosystems, and the causes of this variation are not well understood. Here, we review and synthesize the mechanisms behind CND in fresh waters. We reviewed 131 articles on CND published between 1973 and 1 June 2015. The rate of new publications in CND has increased from 1.4 papers per year during 1973–2002 to 7.3 per year during 2003–2015. The majority of investigations are in North America with many concentrating on fish. More recent studies have focused on animal‐mediated nutrient excretion rates relative to nutrient demand and indirect impacts (e.g. decomposition). We identified several mechanisms that influence CND across levels of biological organization. Factors affecting the stoichiometric plasticity of consumers, including body size, feeding history and ontogeny, play an important role in determining the impact of individual consumers on nutrient dynamics and underlie the stoichiometry of CND across time and space. The abiotic characteristics of an ecosystem affect the net impact of consumers on ecosystem processes by influencing consumer metabolic processes (e.g. consumption and excretion/egestion rates), non‐CND supply of nutrients and ecosystem nutrient demand. Furthermore, the transformation and transport of elements by populations and communities of consumers also influences the flow of energy and nutrients across ecosystem boundaries. This review highlights that shifts in community composition or biomass of consumers and eco‐evolutionary underpinnings can have strong effects on the functional role of consumers in ecosystem processes, yet these are relatively unexplored aspects of CND. Future research should evaluate the value of using species traits and abiotic conditions to predict and understand the effects of consumers on ecosystem‐level nutrient dynamics across temporal and spatial scales. Moreover, new work in CND should strive to integrate knowledge from disparate fields of ecology and environmental science, such as physiology and ecosystem ecology, to develop a comprehensive and mechanistic understanding of the functional role of consumers. Comparative and experimental studies that develop testable hypotheses to challenge the current assumptions of CND, including consumer stoichiometric homeostasis, are needed to assess the significance of CND among species and across freshwater ecosystems.
The metabolic theory of ecology (MTE) and ecological stoichiometry (ES) are both prominent frameworks for understanding energy and nutrient budgets of organisms. We tested their separate and joint ...power to predict nitrogen (N) and phosphorus (P) excretion rates of ectothermic aquatic invertebrate and vertebrate animals (10,534 observations worldwide). MTE variables (body size, temperature) performed better than ES variables (trophic guild, vertebrate classification, body N:P) in predicting excretion rates, but the best models included variables from both frameworks. Size scaling coefficients were significantly lower than predicted by MTE (<0.75), were lower for P than N, and varied greatly among species. Contrary to expectations under ES, vertebrates excreted both N and P at higher rates than invertebrates despite having more nutrient-rich bodies, and primary consumers excreted as much nutrients as carnivores despite having nutrient-poor diets. Accounting for body N:P hardly improved upon predictions from treating vertebrate classification categorically. We conclude that basic data on body size, water temperature, trophic guild, and vertebrate classification are sufficient to make general estimates of nutrient excretion rates for any animal taxon or aquatic ecosystem. Nonetheless, dramatic interspecific variation in size-scaling coefficients and counter-intuitive patterns with respect to diet and body composition underscore the need for field data on consumption and egestion rates. Together, MTE and ES provide a powerful conceptual basis for interpreting and predicting nutrient recycling rates of aquatic animals worldwide.
Nitrogen (N) and phosphorus (P) inputs influence algal community structure and function. The rates and ratios of N and P supply, and different N forms (e.g., NO₃ and NH₄), from external loading and ...internal cycling can be highly seasonal. However, the interaction between seasonality in nutrient supply and algal nutrient limitation remains poorly understood. We examined seasonal variation in nutrient limitation and response to N form in a hypereutrophic reservoir that experiences elevated, but seasonal, nutrient inputs and ratios. External N and P loading is high in spring and declines in summer, when internal loading because more important, reducing loading N:P ratios. Watershed NO₃ dominates spring N supply, but internal NH₄ supply becomes important during summer.We quantified how phytoplankton groups (diatoms, chlorophytes, and cyanobacteria) are limited by N or P, and their N form preference (NH₄ vs. NO₃), with weekly experiments (May–October). Phytoplankton were P-limited in spring, transitioned to N limitation or colimitation (primary N) in summer, and returned to P limitation following fall turnover. UnderNlimitation (or colimitation), chlorophytes and cyanobacteriaweremore strongly stimulated by NH₄ whereas diatoms were often equally, or more strongly, stimulated by NO₃ addition. Cyanobacteria heterocyte development followed the onset of N-limiting conditions,with a severalweek lag time, but heterocyte production did not fully alleviate N-limitation. We show that phytoplankton groups vary seasonally in limiting nutrient and N formpreference, suggesting that dual nutrient management strategies incorporating both N and P, and N formare needed to manage eutrophication.
There is ample evidence that tube-dwelling invertebrates such as chironomids significantly alter multiple important ecosystem functions, particularly in shallow lakes. Chironomids pump large water ...volumes, and associated suspended and dissolved substances, through the sediment and thereby compete with pelagic filter feeders for particulate organic matter. This can exert a high grazing pressure on phytoplankton, microorganisms, and perhaps small zooplankton and thus strengthen benthic-pelagic coupling. Furthermore, intermittent pumping by tube-dwelling invertebrates oxygenates sediments and creates a dynamic, three-dimensional mosaic of redox conditions. This shapes microbial community composition and spatial distribution, and alters microbe-mediated biogeochemical functions, which often depend on redox potential. As a result, extended hotspots of element cycling occur at the oxic-anoxic interfaces, controlling the fate of organic matter and nutrients as well as fluxes of nutrients between sediments and water. Surprisingly, the mechanisms and magnitude of interactions mediated by these organisms are still poorly understood. To provide a synthesis of the importance of tube-dwelling invertebrates, we review existing research and integrate previously disregarded functional traits into an ecosystem model. Based on existing research and our models, we conclude that tube-dwelling invertebrates play a central role in controlling water column nutrient pools, and hence water quality and trophic state. Furthermore, these tiny ecosystem engineers can influence the thresholds that determine shifts between alternate clear and turbid states of shallow lakes. The large effects stand in contrast to the conventional limnological paradigm emphasizing predominantly pelagic food webs. Given the vast number of shallow lakes worldwide, benthic invertebrates are likely to be relevant drivers of biogeochemical processes at regional and global scales, thereby mediating feedback mechanisms linked to climate change.
Climate-change models predict more frequent and intense summer droughts for many areas, including the midwestern United States. Precipitation quantity and intensity in turn drive the rates and ratios ...at which nitrogen (N) and phosphorus (P) are exported from watersheds into lakes, but these rates and ratios are also modulated by watershed land use. This led us to ask the question, is the effect of precipitation on phytoplankton nutrient limitation dependent on watershed land use? Across 42 lakes, we found that phytoplankton in lakes in agricultural landscapes were usually P limited but shifted to strong N limitation under increased drought intensity, and that droughts promoted N-fixing cyanobacteria. In contrast, phytoplankton in lakes with forested watersheds were consistently N limited, regardless of drought status. This climate-land use interaction suggests that droughts may increase the incidence of N limitation in agriculturally impacted lakes. N limitation would likely impair valuable ecosystem services such as drinking water, fisheries, and recreation by promoting the occurrence and severity of cyanobacterial blooms.
Globally, conversion of pristine areas to anthropogenic landscapes is one of the main causes of ecosystem service losses. Land uses associated with urbanization and farming can be major sources of ...pollution to freshwaters promoting artificial inputs of several elements, leading to impaired water quality. However, how the effects of land use on freshwater quality are contingent on properties of the local landscape and climate is still poorly understood. The aim of this study was to evaluate the effects of landscape properties (morphometric measurements of lakes and their catchments), precipitation patterns, and land use properties (extent and proximity of the land use to freshwaters) on water quality of 98 natural lakes and reservoirs in northeast Brazil. Water quality impairment (WQI) was expressed as a composite variable incorporating parameters correlated with eutrophication including nitrogen (N), phosphorus (P) and Chlorophyll-a concentration. Regression tree analysis showed that WQI is mainly related to highly impacted “buffer areas”. However, the effects of land use in these adjacent lands were contingent on precipitation variability for 13% of waterbodies and on surface area of the buffer in relation to the volume of waterbody (BA:Vol) for 87% of waterbodies. Overall, effects on WQI originating from the land use in the adjacent portion of the lake were amplified by high precipitation variability for ecosystems with highly impacted buffer areas and by high BA:Vol for ecosystems with less impacted buffer areas, indicating that ecosystems subjected to intense episodic rainfall events (e.g. storms) and higher buffer areas relative to aquatic ecosystem size (i.e. small waterbodies) are more susceptible to impacts of land use. Land use at the catchment scale was important for the largest ecosystems. Thus, our findings point toward the need for considering a holistic approach to managing water quality, which includes watershed management within the context of climate change.
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•Landscape features and precipitation can mediate land use effects on water quality.•Human land use near shores of lakes and reservoirs decreases their water quality.•Precipitation patterns mediate the effects of land use on water quality.•Climate and geomorphology of lakes are needed to understand land use effects.•Effective policies for climate change and land use are vital to protect freshwater.
Animals can be important in nutrient cycling through a variety of direct and indirect pathways. A high biomass of animals often represents a large pool of nutrients, leading some ecologists to argue ...that animal assemblages can represent nutrient sinks within ecosystems. The role of animals as sources vs. sinks of nutrients has been debated particularly extensively for freshwater fishes. We argue that a large pool size does not equate to a nutrient sink; rather, animals can be nutrient sinks when their biomass increases, when emigration rates are high, and/or when nutrients in animal carcasses are not remineralized. To further explore these ideas, we use a simple model to evaluate the conditions under which fish are phosphorus (P) sources or sinks at the ecosystem (lake) level, and at the habitat level (benthic and water column habitats). Our simulations suggest that, under most conditions, fish are sinks for benthic P but are net P sources to the water column. However, P source and sink strengths depend on fish feeding habits (proportion of P consumed from the benthos and water column), migration patterns, and especially the fate of carcass P. Of particular importance is the rate at which carcasses are mineralized and the relative importance of benthic vs. pelagic primary producers in taking up mineralized P (and excreted P). Higher proportional uptake of P by benthic primary producers increases the likelihood that fish are sinks for water column P. Carcass bones and scales are relatively recalcitrant and can represent a P sink even if fish biomass does not change over time. Thus, there is a need for better documentation of the fraction of carcass P that is remineralized, and the fate of this P, under natural conditions. We urge a more holistic perspective regarding the role of animals in nutrient cycling, with a focus on quantifying the rates at which animals consume, store, release, and transport nutrients under various conditions.
Rates of biogeochemical processes often vary widely in space and time, and characterizing this variation is critical for understanding ecosystem functioning. In streams, spatial hotspots of nutrient ...transformations are generally attributed to physical and microbial processes. Here we examine the potential for heterogeneous distributions of fish to generate hotspots of nutrient recycling. We measured nitrogen (N) and phosphorus (P) excretion rates of 47 species of fish in an N-limited Neotropical stream, and we combined these data with population densities in each of 49 stream channel units to estimate unit- and reach-scale nutrient recycling. Species varied widely in rates of N and P excretion as well as excreted N:P ratios (6-176 molar). At the reach scale, fish excretion could meet >75% of ecosystem demand for dissolved inorganic N and turn over the ambient NH4 pool in <0.3 km. Areal N excretion estimates varied 47-fold among channel units, suggesting that fish distributions could influence local N availability. P excretion rates varied 14-fold among units but were low relative to ambient concentrations. Spatial variation in aggregate nutrient excretion by fish reflected the effects of habitat characteristics (depth, water velocity) on community structure (body size, density, species composition), and the preference of large-bodied species for deep runs was particularly important. We conclude that the spatial distribution of fish could indeed create hotspots of nutrient recycling during the dry season in this species-rich tropical stream. The prevalence of patchy distributions of stream fish and invertebrates suggests that hotspots of consumer nutrient recycling may often occur in stream ecosystems.
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