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      Integrating Stomach Content and Stable Isotope Analyses to Quantify the Diets of Pygoscelid Penguins

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          Abstract

          Stomach content analysis (SCA) and more recently stable isotope analysis (SIA) integrated with isotopic mixing models have become common methods for dietary studies and provide insight into the foraging ecology of seabirds. However, both methods have drawbacks and biases that may result in difficulties in quantifying inter-annual and species-specific differences in diets. We used these two methods to simultaneously quantify the chick-rearing diet of Chinstrap ( Pygoscelis antarctica) and Gentoo ( P. papua) penguins and highlight methods of integrating SCA data to increase accuracy of diet composition estimates using SIA. SCA biomass estimates were highly variable and underestimated the importance of soft-bodied prey such as fish. Two-source, isotopic mixing model predictions were less variable and identified inter-annual and species-specific differences in the relative amounts of fish and krill in penguin diets not readily apparent using SCA. In contrast, multi-source isotopic mixing models had difficulty estimating the dietary contribution of fish species occupying similar trophic levels without refinement using SCA-derived otolith data. Overall, our ability to track inter-annual and species-specific differences in penguin diets using SIA was enhanced by integrating SCA data to isotopic mixing modes in three ways: 1) selecting appropriate prey sources, 2) weighting combinations of isotopically similar prey in two-source mixing models and 3) refining predicted contributions of isotopically similar prey in multi-source models.

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          Most cited references112

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          Combining sources in stable isotope mixing models: alternative methods.

          Stable isotope mixing models are often used to quantify source contributions to a mixture. Examples include pollution source identification; trophic web studies; analysis of water sources for soils, plants; or water bodies, and many others. A common problem is having too many sources to allow a unique solution. We discuss two alternative procedures for addressing this problem. One option is a priori to combine sources with similar signatures so the number of sources is small enough to provide a unique solution. Aggregation should be considered only when isotopic signatures of clustered sources are not significantly different, and sources are related so the combined source group has some functional significance. For example, in a food web analysis, lumping several species within a trophic guild allows more interpretable results than lumping disparate food sources, even if they have similar isotopic signatures. One result of combining mixing model sources is increased uncertainty of the combined end-member isotopic signatures and consequently the source contribution estimates; this effect can be quantified using the IsoError model (http://www.epa.gov/wed/pages/models/isotopes/isoerror1_04.htm). As an alternative to lumping sources before a mixing analysis, the IsoSource mixing model (http://www.epa.gov/wed/pages/models/isosource/isosource.htm) can be used to find all feasible solutions of source contributions consistent with isotopic mass balance. While ranges of feasible contributions for each individual source can often be quite broad, contributions from functionally related groups of sources can be summed a posteriori, producing a range of solutions for the aggregate source that may be considerably narrower. A paleo-human dietary analysis example illustrates this method, which involves a terrestrial meat food source, a combination of three terrestrial plant foods, and a combination of three marine foods. In this case, a posteriori aggregation of sources allowed strong conclusions about temporal shifts in marine versus terrestrial diets that would not have otherwise been discerned.
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            Recent Bayesian stable-isotope mixing models are highly sensitive to variation in discrimination factors.

            Stable isotopes are now used widely in ecological studies, including diet reconstruction, where quantitative inferences about diet composition are derived from the use of mixing models. Recent Bayesian models (MixSIR, SIAR) allow users to incorporate variability in discrimination factors (delta13C or delta15N), or the amount of change in either delta13C or delta15N between prey and consumer, but to date there has been no systematic assessment of the effect of variation in delta13C or delta15N on model outputs. We used whole blood from Common Terns (Sterna hirundo) and muscle from their common prey items (fish and euphausiids) to build a series of mixing models in SIAR (stable isotope analysis in R) using various discrimination factors from the published literature for marine birds. The estimated proportion of each diet component was affected significantly by delta13C or delta15N. We also use recently published stable-isotope data on the reliance of critically endangered Balearic Shearwaters (Puffinus mauretanicus) on fisheries discards to show that discrimination factor choice can have profound implications for conservation and management actions. It is therefore crucial for researchers wishing to use mixing models to have an accurate estimate of delta13C and delta15N, because quantitative diet estimates can help to direct future research or prioritize conservation and management actions.
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              Stable isotopes reveal individual variation in migration strategies and habitat preferences in a suite of seabirds during the nonbreeding period.

              Information on predator and prey distributions is integral to our understanding of migratory connectivity, food web dynamics and ecosystem structure. In marine systems, although large animals that return to land can be fitted with tracking devices, minimum instrument sizes preclude deployments on small seabirds that may nevertheless be highly abundant and hence major consumers. An increasingly popular approach is to use N and C stable isotope analysis of feathers sampled at colonies to provide information on distribution and trophic level for the preceding, and generally little-known, nonbreeding period. Despite the burgeoning of this research, there have been few attempts to verify such relationships. In this study, we demonstrate a clear correspondence between isotope ratios of feathers and nonbreeding distributions of seven species from South Georgia tracked using loggers. This generated a rudimentary isoscape that was used to infer the habitat preferences of eight other species ranging in size from storm petrels to albatrosses, and which could be applied, with caveats, in other studies. Differences in inferred distribution within and between species had major implications for relative exposure to anthropogenic threats, including climate change and fisheries. Although there were no differences in isotope values between sexes in any of the smaller petrels, mean stable C (delta(13)C), but not stable N isotope ratios (delta(15)N), tended to be greater in females than males of the larger, and more sexually size-dimorphic species. This indicates a difference in C source (distribution), rather than trophic level, and a correspondence between the degree of sexual size dimorphism in Procellariiformes and the level of between-sex niche segregation.
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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
                2011
                28 October 2011
                : 6
                : 10
                : e26642
                Affiliations
                [1 ]Department of Biology and Marine Biology, University of North Carolina Wilmington, Wilmington, North Carolina, United States of America
                [2 ]Antarctic Ecosystem Research Division, National Marine Fisheries Service, Southwest Fisheries Science Center, La Jolla, California, United States of America
                [3 ]Department of Biology, Pomona College, Claremont, California, United States of America
                [4 ]Isotope Laboratory, Department of Geological Sciences, Saskatchewan, University of Saskatchewan, Saskatoon, Saskatchewan, Canada
                Phillip Island Nature Parks, Australia
                Author notes

                Conceived and designed the experiments: MJP WZT SDE. Performed the experiments: MJP WZT EN NJK. Analyzed the data: MJP WZT NJK WPP. Wrote the paper: MJP NJK WPP SDE.

                Article
                PONE-D-11-10014
                10.1371/journal.pone.0026642
                3203888
                22053199
                c710a3ed-0ec5-47eb-9356-c4941d253ebd
                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
                : 2 June 2011
                : 30 September 2011
                Page count
                Pages: 10
                Categories
                Research Article
                Biology
                Ecology
                Community Ecology
                Food Web Structure
                Trophic Interactions
                Bioindicators
                Marine Ecology
                Marine Biology
                Marine Ecology
                Zoology
                Ornithology
                Earth Sciences
                Marine and Aquatic Sciences
                Oceans
                Antarctic Ocean
                Marine Biology
                Marine Ecology

                Uncategorized
                Uncategorized

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