Abstract. Local habitat size has been shown to influence colonization and extinction processes of species in patchy environments. However, species differ in body size, mobility, and trophic level, and may not respond in the same way to habitat size. Thus far, we have a limited understanding of how habitat size influences the structure of multitrophic communities and to what extent the effects may be generalizable over a broad geographic range. Here, we used water-filled bromeliads of different sizes as a natural model system to examine the effects of habitat size on the trophic structure of their inhabiting invertebrate communities. We collected composition and biomass data from 651 bromeliad communities from eight sites across Central and South America differing in environmental conditions, species pools, and the presence of large-bodied odonate predators. We found that trophic structure in the communities changed dramatically with changes in habitat (bromeliad) size. Detritivore : resource ratios showed a consistent negative relationship with habitat size across sites. In contrast, changes in predator : detritivore (prey) ratios depended on the presence of odonates as dominant predators in the regional pool. At sites without odonates, predator : detritivore biomass ratios decreased with increasing habitat size. At sites with odonates, we found odonates to be more frequently present in large than in small bromeliads, and predator : detritivore biomass ratios increased with increasing habitat size to the point where some trophic pyramids became inverted. Our results show that the distribution of biomass amongst food-web levels depends strongly on habitat size, largely irrespective of geographic differences in environmental conditions or detritivore species compositions. However, the presence of largebodied predators in the regional species pool may fundamentally alter this relationship between habitat size and trophic structure. We conclude that taking into account the response and multitrophic effects of dominant, mobile species may be critical when predicting changes in community structure along a habitat-size gradient.

Although predators can affect foraging behaviors of floral visitors, rarely is it known if these top-down effects of predators may cascade to plant fitness through trait-mediated interactions. In this study we manipulated artificial crab spiders on flowers of Rubus rosifolius to test the effects of predation risk on flower-visiting insects and strength of trait-mediated indirect effects to plant fitness. In addition, we tested which predator traits (e.g., forelimbs, abdomen) are recognized and avoided by pollinators. Total visitation rate was higher for control flowers than for flowers with an artificial crab spider. In addition, flowers with a sphere (simulating a spider abdomen) were more frequently visited than those with forelimbs or the entire spider model. Furthermore, the presence of artificial spiders decreased individual seed set by 42% and fruit biomass by 50%. Our findings indicate that pollinators, mostly bees, recognize and avoid flowers with predation risk; forelimbs seem to be the predator trait recognized and avoided by hymenopterans. Additionally, predator avoidance by pollinators resulted in pollen limitation, thereby affecting some components of plant fitness (fruit biomass and seed number). Because most pollinator species that recognized predation risk visited many other plant species, trait-mediated indirect effects of spiders cascading down to plant fitness may be a common phenomenon in the Atlantic rainforest ecosystem.

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.

Habitat size and climate are known to affect the trophic structure and dynamics of communities, but their interactive effects are poorly understood. Organisms from different trophic levels vary in terms of metabolic requirements and heat dissipation. Indeed, larger species such as keystone predators require more stable climatic conditions than their prey. Likewise, habitat size disproportionally affects large-sized predators, which require larger home ranges and are thus restricted to larger habitats. Therefore, food web structure in patchy ecosystems is expected to be shaped by habitat size and climate variations. Here we investigate this prediction using natural aquatic microcosm (bromeliad phytotelmata) food webs composed of litter resources (mainly detritus), detritivores, mesopredators, and top predators (damselflies). We surveyed 240 bromeliads of varying sizes (water retention capacity) across 12 open restingas in SE Brazil spread across a wide range of tropical latitudes (-12.6° to -27.6°, ca. 2,000 km) and climates (Δ mean annual temperature = 5.3°C). We found a strong increase in predator-to-detritivore mass ratio with habitat size, which was representative of a typical inverted trophic pyramid in larger ecosystems. However, this relationship was contingent among the restingas; slopes of linear models were steeper in more stable and favorable climates, leading to inverted trophic pyramids (and top-down control) being more pronounced in environments with more favorable climatic conditions. By contrast, detritivore-resource and mesopredator-detritivore mass ratios were not affected by habitat size or climate variations across latitudes. Our results highlight that the combined effects of habitat size, climate and predator composition are pivotal to understanding the impacts of multiple environmental factors on food web structure and dynamics.

Changes in global and regional precipitation regimes are among the most pervasive components of climate change. Intensification of rainfall cycles, ranging from frequent downpours to severe droughts, could cause widespread, but largely unknown, alterations to trophic structure and ecosystem function. We conducted multi-site coordinated experiments to show how variation in the quantity and evenness of rainfall modulates trophic structure in 210 natural freshwater microcosms (tank bromeliads) across Central and South America (18°N to 29°S). The biomass of smaller organisms (detritivores) was higher under more stable hydrological conditions. Conversely, the biomass of predators was highest when rainfall was uneven, resulting in top-heavy biomass pyramids. These results illustrate how extremes of precipitation, resulting in localized droughts or flooding, can erode the base of freshwater food webs, with negative implications for the stability of trophic dynamics.