Environmental heterogeneity at two spatial scales affects litter diversity-decomposition relationships.

The effects of biodiversity on ecological processes have been experimentally evaluated mainly at the local scale under homogeneous conditions. To scale up experimentally based biodiversity-functioning relationships, there is an urgent need to understand how such relationships are affected by the environmental heterogeneity that characterizes larger spatial scales. Here, we tested the effects of an 800-m elevation gradient (a large-scale environmental factor) and forest habitat (a fine-scale factor) on litter diversity-decomposition relationships. To better understand local and landscape scale mechanisms, we partitioned net biodiversity effects into complementarity, selection, and insurance effects as applicable at each scale. We assembled different litter mixtures in aquatic microcosms that simulated natural tree holes, replicating mixtures across blocks nested within forest habitats (edge, interior) and elevations (low, mid, high). We found that net biodiversity and complementarity effects increased over the elevation gradient, with their strength modified by forest habitat and the identity of litter in mixtures. Complementarity effects at local and landscape scales were greatest for combinations of nutrient-rich and nutrient-poor litters, consistent with nutrient transfer mechanisms. By contrast, selection effects were consistently weak and negative at both scales. Selection effects at the landscape level were due mainly to nonrandom overyielding rather than spatial insurance effects. Our findings demonstrate that the mechanisms by which litter diversity affects decomposition are sensitive to environmental heterogeneity at multiple scales. This has implications for the scaling of biodiversity-ecosystem function relationships and suggests that future shifts in environmental conditions due to climate change or land use may impact the functioning of aquatic ecosystems.

Trophic cascades within and across ecosystems: The role of anti-predatory defences, predator type and detritus quality.

Species in one ecosystem can indirectly affect multiple biodiversity components and ecosystem functions of adjacent ecosystems. The magnitude of these cross-ecosystem effects depends on the attributes of the organisms involved in the interactions, including traits of the predator, prey and basal resource. However, it is unclear how predators with cross-ecosystem habitat interact with predators with single-ecosystem habitat to affect their shared ecosystem. Also, unknown is how such complex top-down effects may be mediated by the anti-predatory traits of prey and quality of the basal resource. We used the aquatic invertebrate food webs in tank bromeliads as a model system to investigate these questions. We manipulated the presence of a strictly aquatic predator (damselfly larvae) and a predator with both terrestrial and aquatic habitats (spider), and examined effects on survival of prey (detritivores grouped by anti-predator defence), detrital decomposition (of two plant species differing in litter quality), nitrogen flux and host plant growth. To evaluate the direct and indirect effects each predator type on multiple detritivore groups and ultimately on multiple ecosystem processes, we used piecewise structural equation models. For each response variable, we isolated the contribution of different detritivore groups to overall effects by comparing alternate model formulations. Alone, damselfly larvae and spiders each directly decreased survival of detritivores and caused multiple indirect negative effects on detritus decomposition, nutrient cycling and host plant growth. However, when predators co-occurred, the spider caused a negative non-consumptive effect on the damselfly larva, diminishing the net direct and indirect top-down effects on the aquatic detritivore community and ecosystem functioning. Both detritivore traits and detritus quality modulated the strength and mechanism of these trophic cascades. Predator interference was mediated by undefended or partially defended detritivores as detritivores with anti-predatory defences evaded consumption by damselfly larvae but not spiders. Predators and detritivores affected ecosystem decomposition and nutrient cycling only in the presence of high-quality detritus, as the low-quality detritus was consumed more by microbes than invertebrates. The complex responses of this system to predators from both recipient and adjacent ecosystems highlight the critical role of maintaining biodiversity components across multiple ecosystems.

As espécies em um ecossistema podem afetar indiretamente múltiplos componentes da biodiversidade e funções ecossistêmicas em ecossistemas adjacentes. A magnitude destes efeitos entre ecossistemas depende dos atributos dos organismos envolvidos nas interações, incluindo características do predador, da presa e do recurso basal. No entanto, não está claro como os predadores com habitat em múltiplos ecossistemas interagem com predadores de um ecossistema único, e como isso afeta o ecossistema partilhado entre eles. Além disso, não se sabe como esses efeitos complexos do tipo top­down podem ser mediados pelas características antipredatórias da presa e pela qualidade do recurso basal. Usamos as teias alimentares de invertebrados aquáticos de bromélias­tanque como um sistema modelo para investigar essas questões. Nós manipulamos a presença de um predador estritamente aquático (larvas de zigópteros) e um predador com habitats terrestre e aquático (aranha), e examinamos os efeitos na sobrevivência de presas (grupos de detritívoros com diferentes estratégias de defesa antipredatória), decomposição de detritos foliares (de duas espécies de plantas diferindo na qualidade foliar), fluxo de nitrogênio e crescimento da planta hospedeira. Para avaliar os efeitos diretos e indiretos de cada tipo de predador em múltiplos grupos de detritívoros e, finalmente, em múltiplos processos ecossistêmicos, utilizamos modelos de equações estruturais por partes (piecewiseSEM). Para cada variável resposta, isolamos a contribuição de diferentes grupos de detritívoros bem como seus efeitos globais, comparando modelos alternativos. Larvas de zigópteros e aranhas diminuíram diretamente a sobrevivência dos detritívoros e causaram múltiplos efeitos negativos indiretos na decomposição de detritos, na ciclagem de nutrientes e no crescimento da planta hospedeira. No entanto, quando os predadores coocorreram, a aranha causou um efeito negativo não consumível na larva de zigóptero, diminuindo os efeitos líquidos, diretos e indiretos, do tipo top­down na comunidade de detritívoros aquáticos e no funcionamento do ecossistema. Tanto os atributos antipredatórios dos detritívoros quanto a qualidade dos detritos modularam a força e o mecanismo dessas cascatas tróficas. A interferência do predador foi mediada por detritívoros indefesos ou com defesa parcial. Entretanto, os detritívoros com defesas antipredatórias escaparam do consumo por larvas de zigópteros, mas não por aranhas. Predadores e detritívoros afetaram a decomposição do ecossistema e a ciclagem de nutrientes apenas na presença de detritos de alta qualidade, uma vez que os detritos de baixa qualidade foram consumidos mais por micróbios do que por invertebrados. As respostas complexas deste sistema aos predadores tanto de ecossistemas receptores quanto adjacentes destacam o papel crítico da manutenção dos componentes da biodiversidade em múltiplos ecossistemas.

Do bromeliads affect the arboreal ant communities on orange trees in northwestern Costa Rica?

Ants often interact with other invertebrates as predators or mutualists. Epiphytic bromeliads provide nesting sites for ants, and could increase ant abundances in the tree canopy. We surveyed ants in the foliage of orange trees that either hosted bromeliads or did not. To determine if observed associations between bromeliads and tree ants were causal, we removed bromeliads from half of the trees, and resurveyed ants six weeks later. Our results show that bromeliad presence is correlated with higher ant abundances and different species of ants on orange trees during the dry season. This increase in ant abundance was driven primarily by Solenopsis ants, which were both numerous and found to facultatively nest in bromeliads. Bromeliad removal did not affect either ant abundance or composition, potentially because this manipulation coincided with the transition from dry to wet season. Other ant species were never encountered nesting in bromeliads, and the abundances of such ants on tree leaves were unaffected by bromeliad presence or removal. Considering the importance of ants in herbivore regulation, our findings suggest that bromeliads-through their association with ants-could indirectly be associated with biological control in agricultural systems.

Climate variability and aridity modulate the role of leaf shelters for arthropods: A global experiment.

Current climate change is disrupting biotic interactions and eroding biodiversity worldwide. However, species sensitive to aridity, high temperatures, and climate variability might find shelter in microclimatic refuges, such as leaf rolls built by arthropods. To explore how the importance of leaf shelters for terrestrial arthropods changes with latitude, elevation, and climate, we conducted a distributed experiment comparing arthropods in leaf rolls versus control leaves across 52 sites along an 11,790 km latitudinal gradient. We then probed the impact of short- versus long-term climatic impacts on roll use, by comparing the relative impact of conditions during the experiment versus average, baseline conditions at the site. Leaf shelters supported larger organisms and higher arthropod biomass and species diversity than non-rolled control leaves. However, the magnitude of the leaf rolls' effect differed between long- and short-term climate conditions, metrics (species richness, biomass, and body size), and trophic groups (predators vs. herbivores). The effect of leaf rolls on predator richness was influenced only by baseline climate, increasing in magnitude in regions experiencing increased long-term aridity, regardless of latitude, elevation, and weather during the experiment. This suggests that shelter use by predators may be innate, and thus, driven by natural selection. In contrast, the effect of leaf rolls on predator biomass and predator body size decreased with increasing temperature, and increased with increasing precipitation, respectively, during the experiment. The magnitude of shelter usage by herbivores increased with the abundance of predators and decreased with increasing temperature during the experiment. Taken together, these results highlight that leaf roll use may have both proximal and ultimate causes. Projected increases in climate variability and aridity are, therefore, likely to increase the importance of biotic refugia in mitigating the effects of climate change on species persistence.

Subtle structures with not-so-subtle functions: A data set of arthropod constructs and their host plants.

The construction of shelters on plants by arthropods might influence other organisms via changes in colonization, community richness, species composition, and functionality. Arthropods, including beetles, caterpillars, sawflies, spiders, and wasps often interact with host plants via the construction of shelters, building a variety of structures such as leaf ties, tents, rolls, and bags; leaf and stem galls, and hollowed out stems. Such constructs might have both an adaptive value in terms of protection (i.e., serve as shelters) but may also exert a strong influence on terrestrial community diversity in the engineered and neighboring hosts via colonization by secondary occupants. Although different traits of the host plant (e.g., physical, chemical, and architectural features) may affect the potential for ecosystem engineering by insects, such effects have been, to a certain degree, overlooked. Further analyses of how plant traits affect the occurrence of shelters may therefore enrich our understanding of the organizing principles of plant-based communities. This data set includes more than 1000 unique records of ecosystem engineering by arthropods, in the form of structures built on plants. All records have been published in the literature, and span both natural structures (91% of the records) and structures artificially created by researchers (9% of the records). The data were gathered between 1932 and 2021, across more than 50 countries and several ecosystems, ranging from polar to tropical zones. In addition to data on host plants and engineers, we aggregated data on the type of constructs and the identity of inquilines using these structures. This data set highlights the importance of these subtle structures for the organization of terrestrial arthropod communities, enabling hypotheses testing in ecological studies addressing ecosystem engineering and facilitation mediated by constructs. There are no copyright restrictions and please cite this paper when using the data in publications.

In situ resistance, not immigration, supports invertebrate community resilience to drought intensification in a Neotropical ecosystem.

While future climate scenarios predict declines in precipitations in many regions of the world, little is known of the mechanisms underlying community resilience to prolonged dry seasons, especially in 'naïve' Neotropical rainforests. Predictions of community resilience to intensifying drought are complicated by the fact that the underlying mechanisms are mediated by species' tolerance and resistance traits, as well as rescue through dispersal from source patches. We examined the contribution of in situ tolerance-resistance and immigration to community resilience, following drought events that ranged from the ambient norm to IPCC scenarios and extreme events. We used rainshelters above rainwater-filled bromeliads of French Guiana to emulate a gradient of drought intensity (from 1 to 3.6 times the current number of consecutive days without rainfall), and we analysed the post-drought dynamics of the taxonomic and functional community structure of aquatic invertebrates to these treatments when immigration is excluded (by netting bromeliads) or permitted (no nets). Drought intensity negatively affected invertebrate community resistance, but had a positive influence on community recovery during the post-drought phase. After droughts of 1 to 1.4 times the current intensities, the overall invertebrate abundance recovered within invertebrate life cycle durations (up to 2 months). Shifts in taxonomic composition were more important after longer droughts, but overall, community composition showed recovery towards baseline states. The non-random patterns of changes in functional community structure indicated that deterministic processes like environmental filtering of traits drive community re-assembly patterns after a drought event. Community resilience mostly relied on in situ tolerance-resistance traits. A rescue effect of immigration after a drought event was weak and mostly apparent under extreme droughts. Under climate change scenarios of drought intensification in Neotropical regions, community and ecosystem resilience could primarily depend on the persistence of suitable habitats and on the resistance traits of species, while metacommunity dynamics could make a minor contribution to ecosystem recovery. Climate change adaptation should thus aim at identifying and preserving local conditions that foster in situ resistance and the buffering effects of habitat features.

Extreme rainfall events alter the trophic structure in bromeliad tanks across the Neotropics.

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.

Biogeographic context dependence of trophic cascade strength in bromeliad food webs.

Ecosystem functions and the biomass of lower trophic levels are frequently controlled by predators. The strength of top-down control in these trophic cascades can be affected by the identity and diversity of predators, prey, and resources, as well as environmental conditions such as temperature, moisture, and nutrient loading, which can all impact interaction strength between trophic levels. Few studies have been able to replicate a complete community over a large geographic area to compare the full trophic cascade in a manipulative experiment. Here, we identify geographic dependency in trophic cascade strength, and the driving factors and specific mechanisms behind it, by combining geographically replicated experiments with a novel approach of community analogues of common garden and transplant experiments. We studied a predator-detritivore-detritus food web in bromeliads in Puerto Rico, Costa Rica, and Brazil. We found that interaction strengths between resources, consumers, and predators were strongly site-specific, but the exact mechanism differed between trophic levels. Large bodied predators created strong interaction strengths between predator and consumer trophic levels, reducing consumer abundance regardless of the geographic location, whereas small-bodied predators created weak interactions with no impact on consumer abundances in any site. In contrast, the interaction strength between consumers and resources varied among sites, depending on the dominant species of leaf detritus. More labile leaf species in Costa Rica created a strong consumer-resource interaction and therefore strong trophic cascade, whereas tougher leaf species in Brazil created a weak consumer-resource interaction, and an overall weaker trophic cascade. Our study highlights the importance of replicating experiments over geographic scales to understand general patterns of ecological processes.

Consumptive effects and mismatch in predator-prey turnover rates cause inversion of biomass pyramids.

The mismatch between the turnover rates of predators and prey is one of the oldest explanations for the existence of inverted trophic pyramids. To date, the hypotheses regarding trophic pyramids have all been based on consumptive trophic links between predators and prey, and the relative contribution of non-consumptive effects is still unknown. In this study, we investigated if the inversion of pyramids in bromeliad ecosystems is driven by (i) a rapid colonization of organisms having short cohort interval production (CPI), and (ii) the prevalence of consumptive or non-consumptive effects of top predators. We used a manipulative experiment to investigate the patterns of prey colonization and to partition the net effects of the dominant predator (damselfly larvae) on biomass pyramids into consumptive (uncaged damselfly larvae) and non-consumptive effects (caged damselfly larvae). Consumptive effects of damselflies strengthened the inversion of trophic pyramids. Non-consumptive effects, however, did not affect the shape of biomass pyramids. Instead, the rapid colonization of organisms with predominantly short CPI sustained the large biomass of top predators found in natural bromeliad ecosystems. Prey colonized bromeliads rapidly, but this high production was never visible as standing stock because damselflies reduce prey densities by more than a magnitude through direct consumption. Our study adds to the growing evidence that there are a variety of possible ways that biomass can be trophically structured. Moreover, we suggest that the strength of biomass pyramids inversion may change with the time of ecological succession as prey communities become more equitable.

A precipitation gradient drives change in macroinvertebrate composition and interactions within bromeliads.

Ecological communities change across spatial and environmental gradients due to (i) changes in species composition, (ii) changes in the frequency or strength of interactions or (iii) changes in the presence of the interactions. Here we use the communities of aquatic invertebrates inhabiting clusters of bromeliad phytotelms along the Brazilian coast as a model system for examining variation in multi-trophic communities. We first document the variation in the species pools of sites across a geographical climate gradient. Using the same sites, we also explored the geographic variation in species interaction strength using a Markov network approach. We found that community composition differed along a gradient of water volume within bromeliads due to the spatial turnover of some species. From the Markov network analysis, we found that the interactions of certain predators differed due to differences in bromeliad water volume. Overall, this study illustrates how a multi-trophic community can change across an environmental gradient through changes in both species and their interactions.

Interactive effects of climate change and biodiversity loss on ecosystem functioning.

Climate change and biodiversity loss are expected to simultaneously affect ecosystems, however research on how each driver mediates the effect of the other has been limited in scope. The multiple stressor framework emphasizes non-additive effects, but biodiversity may also buffer the effects of climate change, and climate change may alter which mechanisms underlie biodiversity-function relationships. Here, we performed an experiment using tank bromeliad ecosystems to test the various ways that rainfall changes and litter diversity may jointly determine ecological processes. Litter diversity and rainfall changes interactively affected multiple functions, but how depends on the process measured. High litter diversity buffered the effects of altered rainfall on detritivore communities, evidence of insurance against impacts of climate change. Altered rainfall affected the mechanisms by which litter diversity influenced decomposition, reducing the importance of complementary attributes of species (complementarity effects), and resulting in an increasing dependence on the maintenance of specific species (dominance effects). Finally, altered rainfall conditions prevented litter diversity from fueling methanogenesis, because such changes in rainfall reduced microbial activity by 58%. Together, these results demonstrate that the effects of climate change and biodiversity loss on ecosystems cannot be understood in isolation and interactions between these stressors can be multifaceted.

Effects of predatory ants within and across ecosystems in bromeliad food webs.

Predation is one of the most fundamental ecological processes affecting biotic communities. Terrestrial predators that live at ecosystem boundaries may alter the diversity of terrestrial organisms, but they may also have cross-ecosystem cascading effects when they feed on organisms with complex life cycles (i.e. organisms that shift from aquatic juvenile stages to terrestrial adult stages) or inhibit female oviposition in the aquatic environment. The predatory ant Odontomachus hastatus establishes its colonies among roots of Vriesea procera, an epiphytic bromeliad species with water-filled tanks that shelters many terrestrial and aquatic organisms. Ants may impact terrestrial communities and deter adult insects from ovipositing in the water of bromeliads via consumptive and non-consumptive effects. Ants do not forage within the aquatic environment; thus, they may be more efficient predators on terrestrial organisms. Therefore, we predict that ants will have stronger effects on terrestrial than aquatic food webs. However, such effects may also be site contingent and depend on the local composition of food webs. To test our hypothesis, we surveyed bromeliads with and without O. hastatus colonies from three different coastal field sites in the Atlantic Forest of southeast Brazil, and quantified the effect of this predatory ant on the composition, density and richness of aquatic and terrestrial metazoans found in these bromeliads. We found that ants changed the composition and reduced the overall density of aquatic and terrestrial metazoans in bromeliad ecosystems. However, effects of ants on species diversity were contingent on site. In general terms, the effects of the ant on aquatic and terrestrial metazoan communities were similar in strength and magnitude. Ants reduced the density of virtually all aquatic functional groups, especially detritivore insects as well as metazoans that reach bromeliads through phoresy on the skin of terrestrial animals (i.e. Ostracoda and Helobdella sp.). Our results suggest that the cross-ecosystem effect of this terrestrial predator on the aquatic metazoans was at least as strong as its within-ecosystem effect on the terrestrial ecosystem, and demonstrates that the same predator can simultaneously initiate cascades in multiple ecosystems.

Rainfall and hydrological stability alter the impact of top predators on food web structure and function.

Climate change will alter the distribution of rainfall, with potential consequences for the hydrological dynamics of aquatic habitats. Hydrological stability can be an important determinant of diversity in temporary aquatic habitats, affecting species persistence and the importance of predation on community dynamics. As such, prey are not only affected by drought-induced mortality but also the risk of predation [a non-consumptive effect (NCE)] and actual consumption by predators [a consumptive effect (CE)]. Climate-induced changes in rainfall may directly, or via altered hydrological stability, affect predator-prey interactions and their cascading effects on the food web, but this has rarely been explored, especially in natural food webs. To address this question, we performed a field experiment using tank bromeliads and their aquatic food web, composed of predatory damselfly larvae, macroinvertebrate prey and bacteria. We manipulated the presence and consumption ability of damselfly larvae under three rainfall scenarios (ambient, few large rainfall events and several small rainfall events), recorded the hydrological dynamics within bromeliads and examined the effects on macroinvertebrate colonization, nutrient cycling and bacterial biomass and turnover. Despite our large perturbations of rainfall, rainfall scenario had no effect on the hydrological dynamics of bromeliads. As a result, macroinvertebrate colonization and nutrient cycling depended on the hydrological stability of bromeliads, with no direct effect of rainfall or predation. In contrast, rainfall scenario determined the direction of the indirect effects of predators on bacteria, driven by both predator CEs and NCEs. These results suggest that rainfall and the hydrological stability of bromeliads had indirect effects on the food web through changes in the CEs and NCEs of predators. We suggest that future studies should consider the importance of the variability in hydrological dynamics among habitats as well as the biological mechanisms underlying the ecological responses to climate change.

Predicted rainfall changes disrupt trophic interactions in a tropical aquatic ecosystem.

Changes in the distribution of rainfall and the occurrence of extreme rain events will alter the size and persistence of aquatic ecosystems. Such alterations may affect the structure of local aquatic communities in terms of species composition, and by altering species interactions. In many aquatic ecosystems, leaf litter sustains detrital food webs and could regulate the responses of communities to changes in rainfall. Few empirical studies have focused on how rainfall changes will affect aquatic communities and none have evaluated if basal resource diversity can increase resistance to such rainfall effects. In this study, we used water-holding terrestrial bromeliads, a tropical aquatic ecosystem, to test how predicted rainfall changes and litter diversity may affect community composition and trophic interactions. We used structural equation modeling to investigate the combined effects of rainfall changes and litter diversity on trophic interactions. We demonstrated that changes in rainfall disrupted trophic relationships, even though there were only minor direct effects on species abundance, richness, and community composition. Litter diversity was not able to reduce the impact of changes in rainfall on trophic interactions. We suggest that changes in rainfall can alter the way in which species interact with each other, decreasing the linkages among trophic groups. Such reductions in biotic interactions under climate change will have critical consequences for the functioning of tropical aquatic ecosystems.

Terrestrial support of aquatic food webs depends on light inputs: a geographically-replicated test using tank bromeliads.

Food webs of freshwater ecosystems can be subsidized by allochthonous resources. However, it is still unknown which environmental factors regulate the relative consumption of allochthonous resources in relation to autochthonous resources. Here, we evaluated the importance of allochthonous resources (litterfall) for the aquatic food webs in Neotropical tank bromeliads, a naturally replicated aquatic microcosm. Aquatic invertebrates were sampled in more than 100 bromeliads within either open or shaded habitats and within five geographically distinct sites located in four different countries. Using stable isotope analyses, we determined that allochthonous sources comprised 74% (±17%) of the food resources of aquatic invertebrates. However, the allochthonous contribution to aquatic invertebrates strongly decreased from shaded to open habitats, as light incidence increased in the tanks. The density of detritus in the tanks had no impact on the importance of allochthonous sources to aquatic invertebrates. This overall pattern held for all invertebrates, irrespective of the taxonomic or functional group to which they belonged. We concluded that, over a broad geographic range, aquatic food webs of tank bromeliads are mostly allochthonous-based, but the relative importance of allochthonous subsidies decreases when light incidence favors autochthonous primary production. These results suggest that, for other freshwater systems, some of the between-study variation in the importance of allochthonous subsidies may similarly be driven by the relative availability of autochthonous resources.

Trophic interactions determine the effects of drought on an aquatic ecosystem.

Species interactions can be important mediators of community and ecosystem responses to environmental stressors. However, we still lack a mechanistic understanding of the indirect ecological effects of stress that arise via altered species interactions. To understand how species interactions will be altered by environmental stressors, we need to know if the species that are vulnerable to such stressors also have large impacts on the ecosystem. As predators often exhibit certain traits that are linked to a high vulnerability to stress (e.g., large body size, long generation time), as well as having large effects on communities (e.g., top-down trophic effects), predators may be particularly likely to mediate ecological effects of environmental stress. Other functional groups, like facilitators, are known to have large impacts on communities, but their vulnerability to perturbations remains undocumented. Here, we use aquatic insect communities in bromeliads to examine the indirect effects of an important stressor (drought) on community and ecosystem responses. In a microcosm experiment, we manipulated predatory and facilitative taxa under a range of experimental droughts, and quantified effects on community structure and ecosystem function. Drought, by adversely affecting the top predator, had indirect cascading effects on the entire food web, altering community composition and decomposition. We identified the likely pathway of how drought cascaded through the food web from the top-down as drought -->predator --> shredder --> decomposition. This stress-induced cascade depended on predators exhibiting both a strong vulnerability to drought and large impacts on prey (especially shredders), as well as shredders exhibiting high functional importance as decomposers.

The effects of food web structure on ecosystem function exceeds those of precipitation.

Ecosystems are being stressed by climate change, but few studies have tested food web responses to changes in precipitation patterns and the consequences to ecosystem function. Fewer still have considered whether results from one geographic region can be applied to other regions, given the degree of community change over large biogeographic gradients. We assembled, in one field site, three types of macroinvertebrate communities within water-filled bromeliads. Two represented food webs containing both a fast filter feeder-microbial and slow detritivore energy channels found in Costa Rica and Puerto Rico, and one represented the structurally simpler food webs in French Guiana, which only contained the fast filter feeder-microbial channel. We manipulated the amount and distribution of rain entering bromeliads and examined how food web structure mediated ecosystem responses to changes in the quantity and temporal distribution of precipitation. Food web structure affected the survival of functional groups in general and ecosystem functions such as decomposition and the production of fine particulate organic matter. Ecosystem processes were more affected by decreased precipitation than were the abundance of micro-organisms and metazoans. In our experiments, the sensitivity of the ecosystem to precipitation change was primarily revealed in the food web dominated by the single filter feeder-microbial channel because other top-down and bottom-up processes were weak or absent. Our results show stronger effects of food web structure than precipitation change per se on the functioning of bromeliad ecosystems. Consequently, we predict that ecosystem function in bromeliads throughout the Americas will be more sensitive to changes in the distribution of species, rather than to the direct effects caused by changes in precipitation.

Dominant predators mediate the impact of habitat size on trophic structure in bromeliad invertebrate communities.

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 large-bodied 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.

Resources alter the structure and increase stochasticity in bromeliad microfauna communities.

Although stochastic and deterministic processes have been found to jointly shape structure of natural communities, the relative importance of both forces may vary across different environmental conditions and across levels of biological organization. We tested the effects of abiotic environmental conditions, altered trophic interactions and dispersal limitation on the structure of aquatic microfauna communities in Costa Rican tank bromeliads. Our approach combined natural gradients in environmental conditions with experimental manipulations of bottom-up interactions (resources), top-down interactions (predators) and dispersal at two spatial scales in the field. We found that resource addition strongly increased the abundance and reduced the richness of microfauna communities. Community composition shifted in a predictable way towards assemblages dominated by flagellates and ciliates but with lower abundance and richness of algae and amoebae. While all functional groups responded strongly and predictably to resource addition, similarity among communities at the species level decreased, suggesting a role of stochasticity in species-level assembly processes. Dispersal limitation did not affect the communities. Since our design excluded potential priority effects we can attribute the differences in community similarity to increased demographic stochasticity of resource-enriched communities related to erratic changes in population sizes of some species. In contrast to resources, predators and environmental conditions had negligible effects on community structure. Our results demonstrate that bromeliad microfauna communities are strongly controlled by bottom-up forces. They further suggest that the relative importance of stochasticity may change with productivity and with the organizational level at which communities are examined.

Ecological determinism increases with organism size.

After much debate, there is an emerging consensus that the composition of many ecological communities is determined both by species traits, as proposed by niche theory, as well as by chance events. A critical question for ecology is, therefore, which attributes of species predict the dominance of deterministic or stochastic processes. We outline two hypotheses by which organism size could determine which processes structure ecological communities, and we test these hypotheses by comparing the community structure in bromeliad phytotelmata of three groups of organisms (bacteria, zooplankton, and macroinvertebrates) that encompass a 10 000-fold gradient in body size, but live in the same habitat. Bacteria had no habitat associations, as would be expected from trait-neutral stochastic processes, but still showed exclusion among species pairs, as would be expected from niche-based processes. Macroinvertebrates had strong habitat and species associations, indicating niche-based processes. Zooplankton, with body size between bacteria and macroinvertebrates, showed intermediate habitat associations. We concluded that a key niche process, habitat filtering, strengthened with organism size, possibly because larger organisms are both less plastic in their fundamental niches and more able to be selective in dispersal. These results suggest that the relative importance of deterministic and stochastic processes may be predictable from organism size.