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      Environmental and Organismal Predictors of Intraspecific Variation in the Stoichiometry of a Neotropical Freshwater Fish

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          Abstract

          The elemental composition of animals, or their organismal stoichiometry, is thought to constrain their contribution to nutrient recycling, their interactions with other animals, and their demographic rates. Factors that affect organismal stoichiometry are generally poorly understood, but likely reflect elemental investments in morphological features and life history traits, acting in concert with the environmental availability of elements. We assessed the relative contribution of organismal traits and environmental variability to the stoichiometry of an insectivorous Neotropical stream fish, Rivulus hartii. We characterized the influence of body size, life history phenotype, stage of maturity, and environmental variability on organismal stoichiometry in 6 streams that differ in a broad suite of environmental variables. The elemental composition of R. hartii was variable, and overlapped with the wide range of elemental composition documented across freshwater fish taxa. Average %P composition was ∼3.2%(±0.6), average %N∼10.7%(±0.9), and average %C∼41.7%(±3.1). Streams were the strongest predictor of organismal stoichiometry, and explained up to 18% of the overall variance. This effect appeared to be largely explained by variability in quality of basal resources such as epilithon N∶P and benthic organic matter C∶N, along with variability in invertebrate standing stocks, an important food source for R. hartii. Organismal traits were weak predictors of organismal stoichiometry in this species, explaining when combined up to 7% of the overall variance in stoichiometry. Body size was significantly and positively correlated with %P, and negatively with N∶P, and C∶P, and life history phenotype was significantly correlated with %C, %P, C∶P and C∶N. Our study suggests that spatial variability in elemental availability is more strongly correlated with organismal stoichiometry than organismal traits, and suggests that the stoichiometry of carnivores may not be completely buffered from environmental variability. We discuss the relevance of these findings to ecological stoichiometry theory.

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          Fish extinctions alter nutrient recycling in tropical freshwaters.

          There is increasing evidence that species extinctions jeopardize the functioning of ecosystems. Overfishing and other human influences are reducing the diversity and abundance of fish worldwide, but the ecosystem-level consequences of these changes have not been assessed quantitatively. Recycling of nutrients is one important ecosystem process that is directly influenced by fish. Fish species vary widely in the rates at which they excrete nitrogen and phosphorus; thus, altering fish communities could affect nutrient recycling. Here, we use extensive field data on nutrient recycling rates and population sizes of fish species in a Neotropical river and Lake Tanganyika, Africa, to evaluate the effects of simulated extinctions on nutrient recycling. In both of these species-rich ecosystems, recycling was dominated by relatively few species, but contributions of individual species differed between nitrogen and phosphorus. Alternative extinction scenarios produced widely divergent patterns. Loss of the species targeted by fishermen led to faster declines in nutrient recycling than extinctions in order of rarity, body size, or trophic position. However, when surviving species were allowed to increase after extinctions, these compensatory responses had strong moderating effects even after losing many species. Our results underscore the complexity of predicting the consequences of extinctions from species-rich animal communities. Nevertheless, the importance of exploited species in nutrient recycling suggests that overfishing could have particularly detrimental effects on ecosystem functioning.
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            Light, nutrients, and food-chain length constrain planktonic energy transfer efficiency across multiple trophic levels.

            The efficiency of energy transfer through food chains [food chain efficiency (FCE)] is an important ecosystem function. It has been hypothesized that FCE across multiple trophic levels is constrained by the efficiency at which herbivores use plant energy, which depends on plant nutritional quality. Furthermore, the number of trophic levels may also constrain FCE, because herbivores are less efficient in using plant production when they are constrained by carnivores. These hypotheses have not been tested experimentally in food chains with 3 or more trophic levels. In a field experiment manipulating light, nutrients, and food-chain length, we show that FCE is constrained by algal food quality and food-chain length. FCE across 3 trophic levels (phytoplankton to carnivorous fish) was highest under low light and high nutrients, where algal quality was best as indicated by taxonomic composition and nutrient stoichiometry. In 3-level systems, FCE was constrained by the efficiency at which both herbivores and carnivores converted food into production; a strong nutrient effect on carnivore efficiency suggests a carryover effect of algal quality across 3 trophic levels. Energy transfer efficiency from algae to herbivores was also higher in 2-level systems (without carnivores) than in 3-level systems. Our results support the hypothesis that FCE is strongly constrained by light, nutrients, and food-chain length and suggest that carryover effects across multiple trophic levels are important. Because many environmental perturbations affect light, nutrients, and food-chain length, and many ecological services are mediated by FCE, it will be important to apply these findings to various ecosystem types.
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              Experimental evidence that phenotypic divergence in predators drives community divergence in prey.

              Studies of adaptive divergence have traditionally focused on the ecological causes of trait diversification, while the ecological consequences of phenotypic divergence remain relatively unexplored. Divergence in predator foraging traits, in particular, has the potential to impact the structure and dynamics of ecological communities. To examine the effects of predator trait divergence on prey communities, we exposed zooplankton communities in lake mesocosms to predation from either anadromous or landlocked (freshwater resident) alewives, which have undergone recent and rapid phenotypic differentiation in foraging traits (gape width, gill raker spacing, and prey size-selectivity). Anadromous alewives, which exploit large prey items, significantly reduced the mean body size, total biomass, species richness, and diversity of crustacean zooplankton relative to landlocked alewives, which exploit smaller prey. The zooplankton responses observed in this experiment are consistent with patterns observed in lakes. This study provides direct evidence that phenotypic divergence in predators, even in its early stages, can play a critical role in determining prey community structure.
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                Author and article information

                Contributors
                Role: Editor
                Journal
                PLoS One
                plos
                plosone
                PLoS ONE
                Public Library of Science (San Francisco, USA )
                1932-6203
                2012
                6 March 2012
                : 7
                : 3
                : e32713
                Affiliations
                [1 ]Department of Ecology and Evolutionary Biology, Cornell University, Ithaca, New York, United States of America
                [2 ]School of Natural Resources, University of Nebraska, Lincoln, Nebraska, United States of America
                [3 ]Department of Biology, Drexel University, Philadelphia, Pennsylvania, United States of America
                [4 ]Department of Biological Science, Florida State University, Tallahassee, Florida, United States of America
                [5 ]Odum School of Ecology, University of Georgia, Athens, Georgia, United States of America
                [6 ]Department of Biology, University of California Riverside, Riverside, California, United States of America
                [7 ]Department of Ecology and Evolutionary Biology, Yale University, New Haven, Connecticut, United States of America
                [8 ]Department of Biology, North Carolina State University, Raleigh, North Carolina, United States of America
                Biodiversity Insitute of Ontario - University of Guelph, Canada
                Author notes

                Conceived and designed the experiments: RES ASF SAT DNR CP JT. Performed the experiments: RES MCM TJK EZ. Analyzed the data: RES JT. Contributed reagents/materials/analysis tools: RES TJL EZ JT JFG MW. Wrote the paper: RES.

                [¤a]

                Current address: Department of Biology, University of Victoria, Victoria, British Columbia, Canada

                [¤b]

                Current address: Institute of Arctic and Alpine Research, University of Colorado, Boulder, Colorado, United States of America

                [¤c]

                Current address: Department of Ecology – IBRAG, State University of Rio de Janeiro (UERJ), Rio de Janeiro, Rio de Janeiro, Brazil

                Article
                PONE-D-11-20265
                10.1371/journal.pone.0032713
                3295771
                22412911
                9605bf2d-b7e7-4222-a435-4fdb82323878
                El-Sabaawi et al. This is an open-access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
                History
                : 11 October 2011
                : 30 January 2012
                Page count
                Pages: 12
                Categories
                Research Article
                Biology
                Ecology
                Community Ecology
                Ecosystems
                Evolutionary Biology
                Zoology

                Uncategorized
                Uncategorized

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