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      Ontogenetic and Among-Individual Variation in Foraging Strategies of Northeast Pacific White Sharks Based on Stable Isotope Analysis

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          Abstract

          There is growing evidence for individuality in dietary preferences and foraging behaviors within populations of various species. This is especially important for apex predators, since they can potentially have wide dietary niches and a large impact on trophic dynamics within ecosystems. We evaluate the diet of an apex predator, the white shark ( Carcharodon carcharias), by measuring the stable carbon and nitrogen isotope composition of vertebral growth bands to create lifetime records for 15 individuals from California. Isotopic variations in white shark diets can reflect within-region differences among prey (most importantly related to trophic level), as well as differences in baseline values among the regions in which sharks forage, and both prey and habitat preferences may shift with age. The magnitude of isotopic variation among sharks in our study (>5‰ for both elements) is too great to be explained solely by geographic differences, and so must reflect differences in prey choice that may vary with sex, size, age and location. Ontogenetic patterns in δ 15N values vary considerably among individuals, and one third of the population fit each of these descriptions: 1) δ 15N values increased throughout life, 2) δ 15N values increased to a plateau at ∼5 years of age, and 3) δ 15N values remained roughly constant values throughout life. Isotopic data for the population span more than one trophic level, and we offer a qualitative evaluation of diet using shark-specific collagen discrimination factors estimated from a 3+ year captive feeding experiment (Δ 13C shark-diet and Δ 15N shark-diet equal 4.2‰ and 2.5‰, respectively). We assess the degree of individuality with a proportional similarity index that distinguishes specialists and generalists. The isotopic variance is partitioned among differences between-individual (48%), within-individuals (40%), and by calendar year of sub-adulthood (12%). Our data reveal substantial ontogenetic and individual dietary variation within a white shark population.

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          The ecology of individuals: incidence and implications of individual specialization.

          Most empirical and theoretical studies of resource use and population dynamics treat conspecific individuals as ecologically equivalent. This simplification is only justified if interindividual niche variation is rare, weak, or has a trivial effect on ecological processes. This article reviews the incidence, degree, causes, and implications of individual-level niche variation to challenge these simplifications. Evidence for individual specialization is available for 93 species distributed across a broad range of taxonomic groups. Although few studies have quantified the degree to which individuals are specialized relative to their population, between-individual variation can sometimes comprise the majority of the population's niche width. The degree of individual specialization varies widely among species and among populations, reflecting a diverse array of physiological, behavioral, and ecological mechanisms that can generate intrapopulation variation. Finally, individual specialization has potentially important ecological, evolutionary, and conservation implications. Theory suggests that niche variation facilitates frequency-dependent interactions that can profoundly affect the population's stability, the amount of intraspecific competition, fitness-function shapes, and the population's capacity to diversify and speciate rapidly. Our collection of case studies suggests that individual specialization is a widespread but underappreciated phenomenon that poses many important but unanswered questions.
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            Cascading effects of the loss of apex predatory sharks from a coastal ocean.

            Impacts of chronic overfishing are evident in population depletions worldwide, yet indirect ecosystem effects induced by predator removal from oceanic food webs remain unpredictable. As abundances of all 11 great sharks that consume other elasmobranchs (rays, skates, and small sharks) fell over the past 35 years, 12 of 14 of these prey species increased in coastal northwest Atlantic ecosystems. Effects of this community restructuring have cascaded downward from the cownose ray, whose enhanced predation on its bay scallop prey was sufficient to terminate a century-long scallop fishery. Analogous top-down effects may be a predictable consequence of eliminating entire functional groups of predators.
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              Isotopic ecology ten years after a call for more laboratory experiments.

              About 10 years ago, reviews of the use of stable isotopes in animal ecology predicted explosive growth in this field and called for laboratory experiments to provide a mechanistic foundation to this growth. They identified four major areas of inquiry: (1) the dynamics of isotopic incorporation, (2) mixing models, (3) the problem of routing, and (4) trophic discrimination factors. Because these areas remain central to isotopic ecology, we use them as organising foci to review the experimental results that isotopic ecologists have collected in the intervening 10 years since the call for laboratory experiments. We also review the models that have been built to explain and organise experimental results in these areas.
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                Author and article information

                Contributors
                Role: Editor
                Journal
                PLoS One
                PLoS ONE
                plos
                plosone
                PLoS ONE
                Public Library of Science (San Francisco, USA )
                1932-6203
                2012
                28 September 2012
                : 7
                : 9
                : e45068
                Affiliations
                [1 ]Department of Earth and Planetary Sciences, University of California Santa Cruz, Santa Cruz, California, United States of America
                [2 ]Western Ecological Research Center, United States Geological Survey, Santa Cruz, California, United States of America
                [3 ]Department of Ecology and Evolutionary Biology, University of California Santa Cruz, Santa Cruz, California, United States of America
                University of Leeds, United Kingdom
                Author notes

                Competing Interests: The authors have declared that no competing interests exist.

                Conceived and designed the experiments: SLK PLK. Performed the experiments: SLK MTT JAE PLK. Analyzed the data: SLK MTT JAE PLK. Contributed reagents/materials/analysis tools: SLK MTT JAE PLK. Wrote the paper: SLK MTT JAE PLK.

                Article
                PONE-D-12-08204
                10.1371/journal.pone.0045068
                3460992
                23028766
                f81fc1eb-d6d6-42ee-8062-6e78e3565d4c
                Copyright @ 2012

                This is an open-access article, free of all copyright, and may be freely reproduced, distributed, transmitted, modified, built upon, or otherwise used by anyone for any lawful purpose. The work is made available under the Creative Commons CC0 public domain dedication.

                History
                : 20 March 2012
                : 14 August 2012
                Page count
                Pages: 11
                Funding
                This study was funded by the National Science Foundation grants NSF-OCE 0345943 and NSF-EAR 1053013 to P. Koch and an Institute of Geophysics and Planetary Physics (IGPP) mini-grant award and the Dr. Earl H. Myers and Ethel M. Myers Oceanographic and Marine Biology Trust Award to S. Kim. The funders had no role in study design, data collection and analysis, decision to publish, or preparation of the manuscript.
                Categories
                Research Article
                Agriculture
                Animal Management
                Animal Nutrition
                Biology
                Ecology
                Community Ecology
                Food Web Structure
                Biogeochemistry
                Marine Ecology
                Evolutionary Biology
                Behavioral Ecology
                Marine Biology
                Marine Ecology
                Zoology
                Ichthyology
                Medicine
                Nutrition
                Veterinary Science
                Animal Management
                Animal Nutrition

                Uncategorized
                Uncategorized

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