The niche concept is essential to understanding how biotic and abiotic factors regulate the abundance and distribution of living entities, and how these organisms utilize, affect and compete for resources in the environment. However, it has been challenging to determine the number and types of important niche dimensions. By contrast, there is strong mechanistic theory and empirical evidence showing that the elemental composition of living organisms shapes ecological systems, from organismal physiology to food web structure. We propose an approach based on a multidimensional elemental view of the ecological niche. Visualizing the stoichiometric composition of individuals in multivariate space permits quantification of niche dimensions within and across species. This approach expands on previous elemental characterizations of plant niches, and adapts metrics of niche volume, overlap and nestedness previously used to quantify isotopic niches. We demonstrate the applicability of the multidimensional stoichiometric niche using data on carbon, nitrogen, and phosphorus of terrestrial and freshwater communities composed by multiple trophic groups. First, we calculated the stoichiometric niche volumes occupied by terrestrial and freshwater food webs, by trophic groups, by individual species, and by individuals within species, which together give a measure of the extent of stoichiometric diversity within and across levels of organization. Then we evaluated complementarity between these stoichiometric niches, through metrics of overlap and nestedness. Our case study showed that vertebrates, invertebrates, and primary producers do not overlap in their stoichiometric niches, and that large areas of stoichiometric space are unoccupied by organisms. Within invertebrates, niche differences emerged between freshwater and terrestrial food webs, and between herbivores and non-herbivores (detritivores and predators). These niche differences were accompanied by changes in the covariance structure of the three elements, suggesting fundamental shifts in organismal physiology and/or structure. We also demonstrate the sensitivity of results to sample size, and suggest that representative sampling is better than rarefaction in characterizing the stoichiometric niche occupied by food webs. Overall, our approach demonstrates that stoichiometric traits provide a common currency to estimate the dimensionality of stoichiometric niches, and help reduce and rationalize the number of axis required to characterize communities.

Stoichiometric differences among organisms can affect trophic interactions and rates of nutrient cycling within ecosystems. However, we still know little about either the underlying causes of these stoichiometric differences or the consistency of these differences across large geographical extents. Here, we analyse elemental (carbon, nitrogen, phosphorus) composition of 872 aquatic macroinvertebrates (71 species) inhabiting tank bromeliads (n = 140) from five distantly located sites across Central and South America to (i) test phylogenetic, trophic and body size scaling explanations for why organisms differ in elemental composition and (ii) determine whether patterns in elemental composition are universal or context dependent. Taxonomy explained most variance in elemental composition, even though phylogenetic signals were weak and limited to regional spatial extents and to the family level. The highest elemental contents and lowest carbon:nutrient ratios were found in organisms at high trophic levels and with smaller body size, regardless of geographical location. Carnivores may have higher nutrient content and lower carbon:nutrient ratios than their prey, as organisms optimize growth by choosing the most nutrient‐rich resources to consume and then preferentially retain nutrients over carbon in their bodies. Smaller organisms grow proportionally faster than large organisms and so are predicted to have higher nutrient requirements to fuel RNA and protein synthesis. Geography influenced the magnitude, more than the direction, of the ecological and/or phylogenetic effects on elemental composition. Overall, our results show that both ecological (i.e. trophic group) and evolutionary drivers explain among‐taxa variation in the elemental content of invertebrates, whereas intraspecific variation is mainly a function of body size. Our findings also demonstrate that restricting analyses of macroinvertebrate stoichiometry solely to either the local scale or species level affects inferences of the patterns in invertebrate elemental content and their underlying mechanisms. A http://onlinelibrary.wiley.com/doi/10.1111/1365-2435.13197/suppinfo is available for this article.

Functional traits are commonly used in predictive models that link environmental drivers and community structure to ecosystem functioning. A prerequisite is to identify robust sets of continuous axes of trait variation, and to understand the ecological and evolutionary constraints that result in the functional trait space occupied by interacting species. Despite their diversity and role in ecosystem functioning, little is known of the constraints on the functional trait space of invertebrate biotas of entire biogeographic regions. We examined the ecological strategies and constraints underlying the realized trait space of aquatic invertebrates, using data on 12 functional traits of 852 taxa collected in tank bromeliads from Mexico to Argentina. Principal Component Analysis was used to reduce trait dimensionality to significant axes of trait variation, and the proportion of potential trait space that is actually occupied by all taxa was compared to null model expectations. Permutational Analyses of Variance were used to test whether trait combinations were clade‐dependent. The major axes of trait variation represented life‐history strategies optimizing resource use and antipredator adaptations. There was evidence for trophic, habitat, defence and life‐history niche axes. Bromeliad invertebrates only occupied 16%–23% of the potential space within these dimensions, due to greater concentrations than predicted under uniform or normal distributions. Thus, despite high taxonomic diversity, invertebrates only utilized a small number of successful ecological strategies. Empty areas in trait space represented gaps between major phyla that arose from biological innovations, and trait combinations that are unviable in the bromeliad ecosystem. Only a few phylogenetically distant genera were neighbouring in trait space. Trait combinations aggregated taxa by family and then by order, suggesting that niche conservatism was a widespread mechanism in the diversification of ecological strategies. A http://onlinelibrary.wiley.com/doi/10.1111/1365-2435.13141/suppinfo is available for this article.