Within an organism, lipids are depleted in (13)C relative to proteins and carbohydrates (more negative delta(13)C), and variation in lipid content among organisms or among tissue types has the potential to introduce considerable bias into stable isotope analyses that use delta(13)C. Despite the potential for introduced error, there is no consensus on the need to account for lipids in stable isotope analyses. Here we address two questions: (1) If and when is it important to account for the effects of variation in lipid content on delta(13)C? (2) If it is important, which method(s) are reliable and robust for dealing with lipid variation? We evaluated the reliability of direct chemical extraction, which physically removes lipids from samples, and mathematical normalization, which uses the carbon-to-nitrogen (C:N) ratio of a sample to normalize delta(13)C after analysis by measuring the lipid content, the C:N ratio, and the effect of lipid content on delta(13)C (Deltadelta(13)C) of plants and animals with a wide range of lipid contents. For animals, we found strong relationships between C:N and lipid content, between lipid content and Deltadelta(13)C, and between C:N and Deltadelta(13)C. For plants, C:N was not a good predictor of lipid content or Deltadelta(13)C, but we found a strong relationship between carbon content and lipid content, lipid content and Deltadelta(13)C, and between and carbon content and Deltadelta(13)C. Our results indicate that lipid extraction or normalization is most important when lipid content is variable among consumers of interest or between consumers and end members, and when differences in delta(13)C between end members is <10-12 per thousand. The vast majority of studies using natural variation in delta(13)C fall within these criteria. Both direct lipid extraction and mathematical normalization reduce biases in delta(13)C, but mathematical normalization simplifies sample preparation and better preserves the integrity of samples for delta(15)N analysis.
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.
We use stomach contents and stable isotope ratios of predatory fishes, collected over a 10‐year time span from a species‐rich river in Venezuela, to examine potential body‐size–trophic‐position relationships. Mean body size of predator taxa and their prey (determined by stomach content analyses) were significantly correlated, but trophic position of predators (estimated by stable isotope ratios) was not correlated with body size. This reflects no apparent relationship between body size and trophic position among prey taxa. Primary consumer taxa (algivores and detritivores) in this system are characterized by diverse size and morphology, and thus predatory fish of all body sizes and feeding strategies are able to exploit taxa feeding low in the food web. Regardless of relative body size, predators exploit short, productive food chains. For any given food chain within a complex web where predators are larger than their prey, trophic position and body size are necessarily correlated. But in diverse food webs characterized by a broad range of primary consumer body size, apparently there is no relationship between trophic position and body size across all taxa in the web.
In tropical floodplain rivers, communities associated with structurally complex habitats are disassembled and reassembled as aquatic organisms repeatedly colonize new areas in response to gradual but continuous changes in water level. Thus, a neutral model reflecting random colonization and extinction dynamics may be sufficient to predict assemblage patterns at the scale of local habitat patches. If water level fluctuations and associated patch dynamics are sufficiently predictable, however, community assembly on habitat patches also may be influenced by species-specific responses to habitat features and/or species interactions. We experimentally manipulated structural complexity and proximity to source habitat (which influences colonization rate) of simulated rocky patches in the littoral zone of a tropical lowland river and demonstrate significant effects of both factors on species density of fishes and macroinvertebrates. Interspecific variation in vagility significantly affected assemblage response to habitat complexity. In a second experiment, created habitat patches were sampled over time intervals ranging from 1 day to 36 days to examine temporal dynamics of community assembly. A null-model test revealed that assemblage structure became increasingly non-random, concomitant with increasing species density, over time. Community dynamics in newly formed habitat patches appeared to be dominated by dispersal, whereas in older patches, abundances of individual species increasingly were influenced by habitat characteristics. These data suggest that species-specific responses to environmental variation resulted, in part, because of species interactions. We conclude that community assembly in shallow habitats of this tropical lowland river is influenced by physical habitat characteristics, the spatial distribution of habitat patches, and species interactions as habitats are saturated with individuals.
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