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Species invasions can have substantial impacts on native species and ecosystems, with important consequences for biodiversity. How these disturbances drive changes in the trophic structure of native food webs through time is poorly understood. Here, we quantify trophic disruption in freshwater food webs to invasion by an apex fish predator, lake trout, using an extensive stable isotope dataset across a natural gradient of uninvaded and invaded lakes in the northern Rocky Mountains, USA. Lake trout invasion increased fish diet variability (trophic dispersion), displaced native fishes from their reference diets (trophic displacement), and reorganized macroinvertebrate communities, indicating strong food web disruption. Trophic dispersion was greatest 25 to 50 y after colonization and dissipated as food webs stabilized in later stages of invasion (>50 y). For the native apex predator, bull trout, trophic dispersion preceded trophic displacement, leading to their functional loss in late-invasion food webs. Our results demonstrate how invasive species progressively disrupt native food webs via trophic dispersion and displacement, ultimately yielding biological communities strongly divergent from those in uninvaded ecosystems. [ABSTRACT FROM AUTHOR]

The rapid loss of global biodiversity can greatly affect the functioning of above-ground components of ecosystems. However, how such biodiversity losses affect below-ground communities and linkages to soil carbon (C) sequestration is unclear. Here, we describe how losses in plant functional groups ( PFGs) affect soil microbial and nematode communities and net ecosystem exchange ( NEE) in a 4-year removal experiment conducted on the Mongolian plateau, the world's largest remaining natural grassland., Our results demonstrated that the biomasses or abundances of most components of the two below-ground communities (microbes and nematodes) were negatively affected by PFG loss and were positively related to above-ground plant biomass. The removal of dominant PFGs (perennial bunchgrasses and perennial rhizomatous grasses) reduced the biomass or abundance of below-ground community components while removal of less dominant PFGs (perennial forbs and annuals/biennials) did not change or increased the biomass or abundance of below-ground community components., The biomass-based ratio of fungal to bacterial microbes and the number-based ratio of fungal-feeding to bacterial-feeding nematodes decreased with increasing PFG losses. Variation partitioning analyses showed that the identity of PFGs together with above-ground plant biomass explained most of the total variation in soil microbes and that the identity of PFGs and above-ground plant biomass together with nematode food resources explained most of the total variation in soil nematodes. The increase in NEE with PFG loss was mainly explained by decreases in above-ground plant biomass and the ratio of fungi to bacteria., Synthesis. The shift of below-ground communities from a fungal-based to a bacterial-based energy channel as PFG richness decreases indicates that less diverse grassland ecosystems will have lower nutrient retention and hence be more sensitive to land-use or climate change. The dominant effects of above-ground plant biomass and below-ground communities on NEE indicate that PFG loss resulting from land-use or climate change has the potential to reduce C sequestration in semi-arid grassland soils. These findings suggest that predictive models may need to consider the composition of above-ground and below-ground communities in order to accurately simulate the dynamics of CO2 fluxes in terrestrial ecosystems. [ABSTRACT FROM AUTHOR]

Belowground organisms often display a shift in their massabundance scaling relationships due to environmental factors such as soil chemistry and atmospheric deposition. Here we present new empirical data that show strong differences in allometric scaling according to whether the resolution at the local scale is based on a taxonomic or a functional classification, while only slight differences arise according to soil environmental conditions. For the first time, isometry (an inverse 1 : 1 proportion) is recognized in mass-abundance relationships, providing a functional signal for constant biomass distribution in soil biota regardless of discrete trophic levels. Our findings are in contrast to those from aquatic ecosystems, in that higher trophic levels in soil biota are not a direct function of increasing body mass. [ABSTRACT FROM AUTHOR]

Changes in marine plankton communities driven by environmental variability impact the marine food web and global biogeochemical cycles of carbon and other elements. To predict and assess these community shifts and their consequences, ecologists are increasingly investigating how the functional traits of plankton determine their relative fitness along environmental and biological gradients. Laboratory, field and modelling studies are adopting this trait-based approach to map the biogeography of plankton traits that underlies variations in plankton communities. Here, we review progress towards understanding the regulatory roles of several key plankton functional traits, including cell size, N2-fixation and mixotrophy among phytoplankton, and body size, ontogeny and feeding behaviour for zooplankton. The trait biogeographical approach sheds light on what structures plankton communities in the current ocean, as well as under climate change scenarios, and also allows for finer resolution of community function because community trait composition determines the rates of significant processes, including carbon export. Although understanding of trait biogeography is growing, uncertainties remain that stem, in part, from the paucity of observations describing plankton functional traits. Thus, in addition to recommending widespread adoption of the trait-based approach, we advocate for enhanced collection, standardisation and dissemination of plankton functional trait data. [ABSTRACT FROM AUTHOR]

The transport and deposition of anthropogenic nitrogen (N) to downwind ecosystems is significant and can be a dominant source of new N to many watersheds. Bacterially mediated denitrification in lake sediments may ameliorate the effects of N loading by permanently removing such inputs. We measured denitrification in sediments collected from lakes in the Colorado Rocky Mountains (USA) receiving elevated (5-8 kg N ha y) or low (<2 kg N ha y) inputs of atmospheric N deposition. The nitrate (NO) concentration was significantly greater in high-deposition lakes (11.3 μmol l) compared to low-deposition lakes (3.3 μmol l). Background denitrification was positively related to NO concentrations and we estimate that the sampled lakes are capable of removing a significant portion of N inputs via sediment denitrification. We also conducted a dose-response experiment to determine whether chronic N loading has altered sediment denitrification capacity. Under Michaelis-Menten kinetics, the maximum denitrification rate and half-saturation NO concentration did not differ between deposition regions and were 765 μmol N m h and 293 μmol l NO, respectively, for all lakes. We enumerated the abundances of nitrate- and nitrite-reducing bacteria and found no difference between high- and low-deposition lakes. The abundance of these bacteria was related to available light and bulk sediment resources. Our findings support a growing body of evidence that lakes play an important role in N removal and, furthermore, suggest that current levels of N deposition have not altered the abundance of denitrifying bacteria or saturated the capacity for sediment denitrification. [ABSTRACT FROM AUTHOR]

The article focuses on the evolution of biological stoichiometry under global change. Biological stoichiometry depicts the balance of energy and multiple chemical elements in living systems. Biological stoichiometry and the metabolic theory of ecology bring about greater integration of diverse fields of biology is the geometric framework for nutritional ecology. Biological stoichiometry, with its focus on elements and body composition focuses on nutrients and more. Biological stoichiometry has its centre of gravity at the higher levels of populations, communities and ecosystems.

Nutrient availability and herbivory control the biomass of primary producer communities to varying degrees across ecosystems. Ecological theory, individual experiments in many different systems, and system-specific quantitative reviews have suggested that (i) bottom–up control is pervasive but top–down control is more influential in aquatic habitats relative to terrestrial systems and (ii) bottom–up and top–down forces are interdependent, with statistical interactions that synergize or dampen relative influences on producer biomass. We used simple dynamic models to review ecological mechanisms that generate independent vs. interactive responses of community-level biomass. We calibrated these mechanistic predictions with the metrics of factorial meta-analysis and tested their prevalence across freshwater, marine and terrestrial ecosystems with a comprehensive meta-analysis of 191 factorial manipulations of herbivores and nutrients. Our analysis showed that producer community biomass increased with fertilization across all systems, although increases were greatest in freshwater habitats. Herbivore removal generally increased producer biomass in both freshwater and marine systems, but effects were inconsistent on land. With the exception of marine temperate rocky reef systems that showed positive synergism of nutrient enrichment and herbivore removal, experimental studies showed limited support for statistical interactions between nutrient and herbivory treatments on producer biomass. Top–down control of herbivores, compensatory behaviour of multiple herbivore guilds, spatial and temporal heterogeneity of interactions, and herbivore-mediated nutrient recycling may lower the probability of consistent interactive effects on producer biomass. Continuing studies should expand the temporal and spatial scales of experiments, particularly in understudied terrestrial systems; broaden factorial designs to manipulate independently multiple producer resources (e.g. nitrogen, phosphorus, light), multiple herbivore taxa or guilds (e.g. vertebrates and invertebrates) and multiple trophic levels; and – in addition to measuring producer biomass – assess the responses of species diversity, community composition and nutrient status. [ABSTRACT FROM AUTHOR]

The cycles of the key nutrient elements nitrogen (N) and phosphorus (P) have been massively altered by anthropogenic activities. Thus, it is essential to understand how photosynthetic production across diverse ecosystems is, or is not, limited by N and P. Via a large-scale meta-analysis of experimental enrichments, we show that P limitation is equally strong across these major habitats and that N and P limitation are equivalent within both terrestrial and freshwater systems. Furthermore, simultaneous N and P enrichment produces strongly positive synergistic responses in all three environments. Thus, contrary to some prevailing paradigms, freshwater, marine and terrestrial ecosystems are surprisingly similar in terms of N and P limitation. [ABSTRACT FROM AUTHOR]

Consumer and resource control of diversity in plant communities have long been treated as alternative hypotheses. However, experimental and theoretical evidence suggests that herbivores and nutrient resources interactively regulate the number and relative abundance of coexisting plant species. Experiments have yielded divergent and often contradictory responses within and among ecosystems, and no effort has to date reconciled this empirical variation within a general framework. Using data from 274 experiments from marine, freshwater, and terrestrial ecosystems, we present a cross-system analysis of producer diversity responses to local manipulations of resource supply and/or herbivory. Effects of herbivory and fertilization on producer richness differed substantially between systems: (1) herbivores reduced species richness in freshwater but tended to increase richness in terrestrial systems; (ii) fertilization increased richness in freshwater systems but reduced richness on land. Fertilization consistently reduced evenness, whereas herbivores increased evenness only in marine and terrestrial ecosystems. Producer community evenness and ecosystem productivity mediated fertilization and herbivore effects on diversity across ecosystems. Herbivores increased producer richness in more productive habitats and in producer assemblages with low evenness. These same assemblages also showed the strongest reduction in richness with fertilization, whereas fertilization in- creased (and herbivory decreased) richness in producer assemblages with high evenness. Our study indicates that system productivity and producer evenness determine the direction and magnitude of top-down and bottom-up control of diversity and may reconcile divergent empirical results within and among ecosystems. [ABSTRACT FROM AUTHOR]

The mission of the American Society of Naturalists is "to advance and diffuse knowledge of organic evolution and other broad biological principles so as to enhance the conceptual unification of the biological sciences." In this article, I argue that the area of biology least integrated with knowledge of organic evolution is the field of ecosystem ecology, as evidenced by a semiquantitative literature survey of use of terms in the scientific literature. I present an overview of recent theoretical developments and empirical findings in the emerging field of biological stoichiometry (the study of the balance of energy and multiple chemical elements in living systems). These developments hold some promise as a means to conceptually integrate ecosystem ecology, with its emphasis on flows and pools of energy and chemical elements, with evolutionary biology, with its emphasis on genetic fitness and the biochemical products of the genome. For example, recent evidence indicates that organismal C : P and N : P ratios have a major impact on biologically mediated flows of energy and phosphorus; in turn, variations among taxa in these ratios are connected to evolved differences in organismal growth rate because of the connection between growth rate and the need for increased allocation to P-rich ribosomal RNA. In this way, evolutionary change in growth-related traits, by altering organismal P requirements, has direct biogeochemical implications, while ecosystem conditions can constrain evolutionary acceleration of growth rates by imposing a direct P limitation on production of the needed biochemical machinery of growth. Thus, stoichiometric theory provides a broad biological principle that can interconvert the currencies and concerns of ecosystem ecology and evolutionary biology, facilitating integration of diverse fields of study and contributing to conceptual unification of the biological sciences. [ABSTRACT FROM AUTHOR]

Plant biomass and nutrient allocation explicitly links the evolved strategies of plant species to the material and energy cycles of ecosystems. Allocation of nitrogen (N) and phosphorus (P) is of particular interest because N and P play pivotal roles in many aspects of plant biology, and their availability frequently limits plant growth. Here we present a comparative scaling analysis of a global data compilation detailing the N and P contents of leaves, stems, roots, and reproductive structures of 1,287 species in 152 seed plant families. We find that P and N contents (as well as N : P) are generally highly correlated both within and across organs and that differences exist between woody and herbaceous taxa. Between plant organs, the quantitative form of the scaling relationship changes systematically, depending on whether the organs considered are primarily structural (i.e., stems, roots) or metabolically active (i.e., leaves, reproductive structures). While we find significant phylogenetic signals in the data, similar scaling relationships occur in independently evolving plant lineages, which implies that both the contingencies of evolutionary history and some degree of environmental convergence have led to a common set of rules that constrain the partitioning of nutrients among plant organs. [ABSTRACT FROM AUTHOR]

The Cuatro Cienegas basin in the Chihuahuan desert is a system of springs, streams, and pools. These ecosystems support >70 endemic species and abundant living stromatolites and other microbial communities, representing a desert oasis of high biodiversity. Here, we combine data from molecular microbiology and geology to document the microbial biodiversity of this unique environment. Ten water samples from locations within the Cuatro Cienegas basin and two neighboring valleys as well as three samples of wet sediments were analyzed. The phylogeny of prokaryotic populations in the samples was determined by characterizing cultured organisms and by PCR amplification and sequencing of 16S rRNA genes from total community DNA. The composition of microbial communities was also assessed by determining profiles of terminal restriction site polymorphisms of 16S rRNA genes in total community DNA. There were 250 different phylotypes among the 350 cultivated strains. Ninety-eight partial 16S rRNA gene sequences were obtained and classified. The clones represented 38 unique phylotypes from ten major lineages of Bacteria and one of Archaea. Unexpectedly, 50% of the phylotypes were most closely related to marine taxa, even though these environments have not been in contact with the ocean for tens of millions of years. Furthermore, terminal restriction site polymorphism profiles and geological data suggest that the aquatic ecosystems of Cuatro Cienegas are hydrologically interconnected with adjacent valleys recently targeted for agricultural intensification. The findings underscore the conservation value of desert aquatic ecosystems and the urgent need for study and preservation of freshwater microbial communities. [ABSTRACT FROM AUTHOR]

Here we use the theory of ecological stoichiometry to propose and provide a preliminary test of a novel hypothesis that the Cambrian ‘explosion’ may have been triggered by changes in circulating P availability in the biosphere. We exposed living stromatolites from a spring-fed stream in Mexico to a gradient of P enrichment to examine subsequent effects on stromatolite C : P ratio and on the primary grazer, an endemic snail. Consistent with a previously hypothesized stoichiometric ‘knife-edge’, snail performance was maximal at intermediate P-enrichment, indicating in situ stoichiometric constraints because of high stromatolite C : P ratio along with high sensitivity to excessive P intake. These results are consistent with the idea that stoichiometric constraints may have delayed the evolutionary proliferation of animals in ancient stromatolite-dominated ecosystems and also suggest that high food P content can significantly impair consumers. We propose that ecosystem P availability may have impacted both the expansion and decline of animal taxa in the history of life. [ABSTRACT FROM AUTHOR]

Understanding the storage, flux, and turnover of nutrients in organisms is important for quantifying contributions of biota to biogeochemical cycles. Here we present a model that predicts the storage of phosphorus-rich RNA and whole-body phosphorus content in eukaryotes based on the mass- and temperature-dependence of ATP production in mitochondria. Data from a broad assortment of eukaryotes support the model's two main predictions. First, whole-body RNA concentration is proportional to mitochondrial density and consequently scales with body mass to the -1/4 power. Second, whole-body phosphorus content declines with increasing body mass in eukaryotic unicells but approaches a relatively constant value in large multicellular animals because the fraction of phosphorus in RNA decreases relative to the fraction in other pools. Extension of the model shows that differences in the flux of RNA-associated phosphorus are due to the size dependencies of metabolic rate and RNA concentration. Thus, the model explicitly links two biological currencies at the individual level: energy in the form of ATP and materials in the form of phosphorus, both of which are critical to the functioning of ecosystems. The model provides a framework for linking attributes of individuals to the storage and flux of phosphorus in ecosystems. [ABSTRACT FROM AUTHOR]

Animals encountering nutritionally imbalanced foods should release elements in excess of requirements in order to maintain overall homeostasis. Quantifying these excesses and predicting their fate is, however, problematic. A new model of the stoichiometry of consumers is formulated that incorporates the separate terms in the metabolic budget, namely, assimilation of ingested substrates and associated costs, protein turnover, other basal costs, such as osmoregulation, and the use of remaining substrates for production. The model indicates that release of excess C and non limiting nutrients may often be a significant fraction of the total metabolic budget of animals consuming the nutrient-deficient forages that are common in terrestrial and aquatic systems. The cost of maintenance, in terms of not just C but also N and P, is considerable, such that food quality is important even when intake is low. Many generalist consumers experience short-term and unpredictable fluctuations in their diets. Comparison of model output with data for one such consumer, Daphnia, indicates that mechanisms operating postabsorption in the gut are likely the primary means of regulating excess C, N, and P in these organisms, notably respiration decoupled from biochemical or mechanical work and excretion of carbon and nutrients. This stoichiometrically regulated release may often be in organic rather than inorganic form, with important consequences for the balance of autotrophic and heterotrophic processes in ecosystems. [ABSTRACT FROM AUTHOR]

The fate of carbon (C) in organisms, food webs, and ecosystems is to a major extent regulated by mass-balance principles and the availability of other key nutrient elements. In relative terms, nutrient limitation implies excess C, yet the fate of this C may be quite different in autotrophs and heterotrophs. For autotrophs nutrient limitation means less fixation of inorganic C or excretion of organic C, while for heterotrophs nutrient limitation means that more of ingested C will "go to waste" in the form of egestion or respiration. There is in general a mismatch between autotrophs and decomposers that have flexible but generally high C:element ratios, and consumers that have lower C:element ratios and tighter stoichiometric regulation. Thus, C-use efficiency in food webs may be governed by the element ratios in autotroph biomass and tend to increase when C:element ratios in food approach those of consumers. This tendency, has a strong bearing on the sequestration of C in ecosystems, since more C will be diverted to detritus entering soils or sediments when C-use efficiency is low due to stoichiometric imbalance. There will be a strong evolutionary pressure to utilize such excess C for structural and metabolic purposes. This article explores how these basic principles may regulate C sequestration on different scales in aquatic and terrestrial ecosystems. [ABSTRACT FROM AUTHOR]

Understanding the reciprocal interactions between the evolved characteristics of species and the environment in which each species is embedded is a major priority for evolutionary ecology. Here we use the perspective of ecological stoichiometry to test the hypothesis that natural selection on body growth rate affects consumer body stoichiometry. As body elemental composition (nitrogen, phosphorus) of consumers influences nutrient cycling and trophic dynamics in food webs, such differences should also affect biogeochemical processes and trophic dynamics. Consistent with the growth rate hypothesis, body growth rate and phosphorus content of individuals of the Daphnia pulex species complex were lower in Wisconsin compared to Alaska, where the brevity of the growing season places a premium on growth rate. Consistent with stoichiometric theory, we also show that, relative to animals sampled in Wisconsin, animals sampled in Alaska were poor recyclers of P and suffered greater declines in growth when fed low-quality, P-deficient food. These results highlight the importance of evolutionary context in establishing the reciprocal relationships between single species and ecosystem processes such as trophic dynamics and consumer-driven nutrient recycling. [ABSTRACT FROM AUTHOR]

Examines the relative balance of light and phosphorus in a set of freshwater pelagic ecosystems. Methodology used in study; Calculation of a ratio of light; Correlation between the C:P ratio of suspended particulate matter and the light: phosphorus ratio within the epilimnion; Hypothesized trends in ecosystem function; Discussion on study findings; Concluding remarks.

Comments on the elements of ecology and evolution. Ecological stoichiometry as the study of the balance of energy and multiple chemical elements in ecological interactions; Expansion of the concept of ecological stoichiometry to other organism and other ecosystems; Applications and horizons for stoichiometric theory.

1. Cyanobacteria blooms have long been the focus of limnological research as they often represent the ‘end member’ of limnological deterioration under various human impacts. Research over the past several decades has greatly illuminated the ecological factors promoting cyanobacteria blooms but controversy and confusion surround the successful integration of this diverse body of work. 2. In this opinion article I attempt to integrate well-known aspects of cyanobacteria bloom ecology (such as the roles of nutrient loading, N:P ratio, and mixing conditions) with more recent developments that highlight the importance of feedbacks within the food web in regulating cyanobacteria blooms. Food-web feedbacks involving stoichiometric mechanisms appear to be particularly important, as accumulating data indicate that the food web influences cyanobacteria not just by regulating the rate of grazing mortality. Rather, trophic interactions may also regulate cyanobacteria dynamics by altering the consumer-driven nutrient recycling regime in a way that shifts the competitive advantage away from cyanobacteria. 3. Viewed in this way, cyanobacteria blooms can be seen as probabilistic events that are the end result of a series of key mechanisms involving nutrient loading, physical mixing conditions, and trophic interactions. To successfully manage lake water quality we should take advantage of each node of contingency leading to undesirable blooms. In doing so we will also have a more coherent scientific message to communicate with those directly involved with the socioeconomic politics of water quality decision making. [ABSTRACT FROM AUTHOR]

The competitive exclusion principle (CEP) states that no equilibrium is possible if n species exploit fewer than n resources. This principle does not appear to hold in nature, where high biodiversity is commonly observed, even in seemingly homogenous habitats. Although various mechanisms, such as spatial heterogeneity or chaotic fluctuations, have been proposed to explain this coexistence, none of them invalidates this principle. Here we evaluate whether principles of ecological stoichiometry can contribute to the stable maintenance of biodiverse communities. Stoichiometric analysis recognizes that each organism is a mixture of multiple chemical elements such as carbon (C), nitrogen (N), and phosphorus (P) that are present in various proportions in organisms. We incorporate these principles into a standard predator–prey model to analyze competition between two predators on one autotrophic prey. The model tracks two essential elements, C and P, in each species. We show that a stable equilibrium is possible with two predators on this single prey. At this equilibrium both predators can be limited by the P content of the prey. The analysis suggests that chemical heterogeneity within and among species provides new mechanisms that can support species coexistence and that may be important in maintaining biodiversity. [Copyright &y& Elsevier]