Stable isotope ratios (typically of carbon and nitrogen) provide one representation of an organism's trophic niche and are widely used to examine aspects of food web structure. Yet stable isotopes have not been applied to quantitatively characterize community-wide aspects of trophic structure (i.e., at the level of an entire food web). We propose quantitative metrics that can be used to this end, drawing on similar approaches from ecomorphology research. For example, the convex hull area occupied by species in delta13C-delta15N niche space is a representation of the total extent of trophic diversity within a food web, whereas mean nearest neighbor distance among all species pairs is a measure of species packing within trophic niche space. To facilitate discussion of opportunities and limitations of the metrics, we provide empirical and conceptual examples drawn from Bahamian tidal creek food webs. These examples illustrate how this methodology can be used to quantify trophic diversity and trophic redundancy in food webs, as well as to link individual species to characteristics of the food web in which they are embedded. Building from extensive applications of stable isotope ratios by ecologists, the community-wide metrics may provide a new perspective on food web structure, function, and dynamics.

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Can stable isotope ratios provide for community-wide measures of trophic structure?

Fifty years ago, GE Hutchinson defined the ecological niche as a hypervolume in n-dimensional space with environmental variables as axes. Ecologists have recently developed renewed interest in the concept, and technological advances now allow us to use stable isotope analyses to quantify these niche dimensions. Analogously, we define the isotopic niche as an area (in δ-space) with isotopic values (δ-values) as coordinates. To make isotopic measurements comparable to other niche formulations, we propose transforming δ-space to p-space, where axes represent relative proportions of isotopically distinct resources incorporated into an animal's tissues. We illustrate the isotopic niche with two examples: the application of historic ecology to conservation biology and ontogenetic niche shifts. Sustaining renewed interest in the niche requires novel methods to measure the variables that define it. Stable isotope analyses are a natural, perhaps crucial, tool in contemporary studies of the ecological niche.

A niche for isotopic ecology

Stable isotopes are a powerful tool for ecologists, often used to assess contributions of different sources to a mixture (e.g. prey to a consumer). Mixing models use stable isotope data to estimate the contribution of sources to a mixture. Uncertainty associated with mixing models is often substantial, but has not yet been fully incorporated in models. We developed a Bayesian-mixing model that estimates probability distributions of source contributions to a mixture while explicitly accounting for uncertainty associated with multiple sources, fractionation and isotope signatures. This model also allows for optional incorporation of informative prior information in analyses. We demonstrate our model using a predator–prey case study. Accounting for uncertainty in mixing model inputs can change the variability, magnitude and rank order of estimates of prey (source) contributions to the predator (mixture). Isotope mixing models need to fully account for uncertainty in order to accurately estimate source contributions.

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Incorporating uncertainty and prior information into stable isotope mixing models

Studies in recent years have suggested that the conservation of biodiversity improves the ability of an ecosystem to retain nutrients and remain productive. These papers have proved controversial, in part because of a lack of direct evidence for a mechanism to explain the phenomenon. Now, in experiments involving manipulation of the number of algal species in model stream systems, Bradley Cardinale provides one such mechanism. Uptake of nitrogen nutrients increased linearly with species richness in response to changes in flow habitats and disturbance regimes. But when niche structure was experimentally removed, the relationship disappeared. This suggests that habitats with more species take greater advantage of the niche opportunities in an environment than do less-species-rich habitats, allowing the more diverse systems to capture a greater fraction of biologically active resources such as nitrogen. More diverse stream communities have increased uptake of nutrients, including nitrate, a major pollutant, but the mechanism is little understood. This study manipulated algal species diversity in stream mesocosms with different flow habitats and disturbance regimes. Nitrogen uptake increased linearly with species richness, but when niche structure was experimentally removed the relationship disappeared. Excessive nutrient loading of water bodies is a leading cause of water pollution worldwide1,2, and controlling nutrient levels in watersheds is a primary objective of most environmental policy3. Over the past two decades, much research has shown that ecosystems with more species are more efficient at removing nutrients from soil and water than are ecosystems with fewer species4,5,6,7. This has led some to suggest that conservation of biodiversity might be a useful tool for managing nutrient uptake and storage7,8,9,10, but this suggestion has been controversial, in part because the specific biological mechanisms by which species diversity influences nutrient uptake have not been identified10,11,12. Here I use a model system of stream biofilms to show that niche partitioning among species of algae can increase the uptake and storage of nitrate, a nutrient pollutant of global concern. I manipulated the number of species of algae growing in the biofilms of 150 stream mesocosms that had been set up to mimic the variety of flow habitats and disturbance regimes that are typical of natural streams. Nitrogen uptake rates, as measured by using 15N-labelled nitrate, increased linearly with species richness and were driven by niche differences among species. As different forms of algae came to dominate each unique habitat in a stream, the more diverse communities achieved a higher biomass and greater 15N uptake. When these niche opportunities were experimentally removed by making all of the habitats in a stream uniform, diversity did not influence nitrogen uptake, and biofilms collapsed to a single dominant species. These results provide direct evidence that communities with more species take greater advantage of the niche opportunities in an environment, and this allows diverse systems to capture a greater proportion of biologically available resources such as nitrogen. One implication is that biodiversity may help to buffer natural ecosystems against the ecological impacts of nutrient pollution.

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Biodiversity improves water quality through niche partitioning

Stable isotope analysis (SIA) has emerged as a common tool in ecology and has proven especially useful in the study of animal diet, habitat use, movement, and physiology. SIA has been vigorously applied to the study of marine mammals, because most species live in habitats or undergo large migrations/movements that make them difficult to observe. Our review supplies a complete list of published SIA contributions to marine mammal science and highlights informative case examples in four general research areas: (1) physiology and fractionation, (2) foraging ecology and habitat use, (3) ecotoxicology, and (4) historic ecology and paleoecology. We also provide a condensed background of isotopic nomenclature, highlight several physiological considerations important for accurate interpretation of isotopic data, and identify research areas ripe for future growth. Because it is impossible to conduct controlled laboratory experiments on most marine mammal species, future studies in marine mammal ecology must draw on isotopic data collected from other organisms and be cognizant of key assumptions often made in the application of SIA to the study of animal ecology. The review is designed to be accessible to all audiences, from students unfamiliar with SIA to those who have utilized it in published studies.

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Using stable isotope biogeochemistry to study marine mammal ecology

Isotope Notation and Measurement.- Using Stable Isotope Tracers.- Isotope Chi.- Mixing.- Isotope Additions.- Fractionation.- Scanning the Future.

Brian Fry

Papers

Stable isotope ecology