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.
Diverse invertebrate and vertebrate species live in association with plants of the large Neotropical family Bromeliaceae. Although previous studies have assumed that debris of associated organisms improves plant nutrition, so far little evidence supports this assumption. In this study we used isotopic ((15)N) and physiological methods to investigate if the treefrog Scinax hayii, which uses the tank epiphytic bromeliad Vriesea bituminosa as a diurnal shelter, contributes to host plant nutrition. In the field, bromeliads with frogs had higher stable N isotopic composition (delta(15)N) values than those without frogs. Similar results were obtained from a controlled greenhouse experiment. Linear mixing models showed that frog feces and dead termites used to simulate insects that eventually fall inside the bromeliad tank contributed, respectively, 27.7% (+ or - 0.07 SE) and 49.6% (+ or - 0.50 SE) of the total N of V. bituminosa. Net photosynthetic rate was higher in plants that received feces and termites than in controls; however, this effect was only detected in the rainy, but not in the dry season. These results demonstrate for the first time that vertebrates contribute to bromeliad nutrition, and that this benefit is seasonally restricted. Since amphibian-bromeliad associations occur in diverse habitats in South and Central America, this mechanism for deriving nutrients may be important in bromeliad systems throughout the Neotropics.
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These results indicate that the mutualism between spiders and bromeliads is seasonally restricted, generating a conditional outcome. There was interspecific variation in nutrient uptake, probably related to each species' performance and photosynthetic pathways. Whereas B. balansae seems to use nitrogen for growth, Ae. distichantha apparently stores nitrogen for stressful nutritional conditions. Bromeliads absorbed more nitrogen coming from spider faeces than from flies, reinforcing the beneficial role played by predators in these digestive mutualisms.
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