Effects of warming on the structure of aquatic communities in tropical bromeliad microecosystems.

Freshwaters are among the most vulnerable ecosystems to climate warming, with projected temperature increases over the coming decades leading to significant losses of aquatic biodiversity. Experimental studies that directly warm entire natural ecosystems in the tropics are needed, for understanding the disturbances on aquatic communities. Therefore, we conducted an experiment to test the impacts of predicted future warming on density, alpha diversity, and beta diversity of freshwater aquatic communities, inhabiting natural microecosystems-Neotropical tank bromeliads. Aquatic communities within the tanks bromeliads were experimentally exposed to warming, with temperatures ranging from 23.58 to 31.72°C. Linear regression analysis was used to test the impacts of warming. Next, distance-based redundancy analysis was performed to assess how warming might alter total beta diversity and its components. This experiment was conducted across a gradient of habitat size (bromeliad water volume) and availability of detrital basal resources. A combination of the highest detritus biomass and higher experimental temperatures resulted in the greatest density of flagellates. However, the density of flagellates declined in bromeliads with higher water volume and lower detritus biomass. Moreover, the combination of the highest water volume and high temperature reduced density of copepods. Finally, warming changed microfauna species composition, mostly through species substitution (ß repl component of total beta-diversity). These findings indicate that warming strongly structures freshwater communities by reducing or increasing densities of different aquatic communities groups. It also enhances beta-diversity, and many of these effects are modulated by habitat size or detrital resources.

Local drivers of heterogeneity in a tropical forest: epiphytic tank bromeliads affect the availability of soil resources and conditions and indirectly affect the structure of seedling communities.

Environmental heterogeneity is a key component in explaining the megadiversity of tropical forests. Despite its importance, knowledge about local drivers of environmental heterogeneity remains a challenge for ecologists. In Neotropical forests, epiphytic tank bromeliads store large amounts of water and nutrients in the tree canopy, and their tank overflow may create nutrient-rich patches in the soil. However, the effects of this nutrient flux on environmental heterogeneity and plant community structure in the understory remain unexplored. In a Brazilian coastal sandy forest, we investigated the effects of the presence of epiphytic tank bromeliads on throughfall chemistry, soil chemistry, soil litter biomass, light, and seedling community structure. In the presence of epiphytic tank bromeliads, the throughfall nitrogen concentration increased twofold, the throughfall phosphorus concentration increased threefold, and the soil patches had a 3.96% higher pH, a 50% higher calcium concentration, and 11.88% less light. By altering the availability of soil resources and conditions, the presence of bromeliads partially shifted the available niche spaces for plant species and indirectly affected the structure of the seedling communities, decreasing their diversity, density, and biomass. For the first time, we showed that the presence of tank bromeliads in the canopy can create characteristic soil patches in the understory, affecting the structure of seedling communities via fertilization. Our results reveal a novel local driver of environmental heterogeneity, reinforcing and expanding the key role of tank bromeliads both in nutrient cycling and plant community structuring of Neotropical coastal sandy forests.

A neotropical mistletoe influences herbivory of its host plant by driving changes in the associated insect community.

Parasitic plants are important sources of stress and can strongly impact their host plants through direct and indirect associations with other herbivores and their associated organisms. In the tropics, mistletoes are frequent parasitic plants, influencing different trophic levels involved with the host plant. Here, we investigated the direct and indirect influences of multiple partners involved in interactions between the mistletoe Psittachantus robustus and its host tree, Vochysia thyrsoidea. More specifically, we assessed if the presence of the mistletoe modified herbivory levels of its host by altering the diversity of associated insects. We found that insect feeding guild modulated mistletoe influence on insect community, and there were fewer species and individuals of leaf-chewing insects in parasitized than non-parasitized trees. Despite this decrease in leaf-chewing insects, there were increased levels of herbivory in parasitized plants. Mistletoes' presence did not influence the hemipteran sap-sucking insects, but this herbivore guild directly responded to the abundance of their associated ants. Overall, our study found empirical support for the crucial role of mistletoes on their host-associated organisms, ultimately shaping the herbivory levels of their tree hosts. By exposing the distinct effects of the different partners involved, our results shed light on the intricated interactions mediated by parasitic plants, opening the path for new investigations.

Regime shifts in a shallow lake over 12 years: Consequences for taxonomic and functional diversities, and ecosystem multifunctionality.

Under increasing nutrient loading, shallow lakes may shift from a state of clear water dominated by submerged macrophytes to a turbid state dominated by phytoplankton or a shaded state dominated by floating macrophytes. How such regime shifts mediate the relationship between taxonomic and functional diversities (FD) and lake multifunctionality is poorly understood. We employed a detailed database describing a shallow lake over a 12-year period during which the lake has displayed all the three states (clear, turbid and shaded) to investigate how species richness, FD of fish and zooplankton, ecosystem multifunctionality and five individual ecosystem functions (nitrogen and phosphorus concentrations, standing fish biomass, algae production and light availability) differ among states. We also evaluated how the relationship between biodiversity (species richness and FD) and multifunctionality is affected by regime shifts. We showed that species richness and the FD of fish and zooplankton were highest during the clear state. The clear state also maintained the highest values of multifunctionality as well as standing fish biomass production, algae biomass and light availability, whereas the turbid and shaded states had higher nutrient concentrations. Functional diversity was the best predictor of multifunctionality. The relationship between FD and multifunctionality was strongly positive during the clear state, but such relationship became flatter after the shift to the turbid or shaded state. Our findings illustrate that focusing on functional traits may provide a more mechanistic understanding of how regime shifts affect biodiversity and the consequences for ecosystem functioning. Regime shifts towards a turbid or shaded state negatively affect the taxonomic diversity and FD of fish and zooplankton, which in turn impairs the multifunctionality of shallow lakes.

Trophic downgrading decreases species asynchrony and community stability regardless of climate warming.

Theory and some evidence suggest that biodiversity promotes stability. However, evidence of how trophic interactions and environmental changes modulate this relationship in multitrophic communities is lacking. Given the current scenario of biodiversity loss and climate changes, where top predators are disproportionately more affected, filling these knowledge gaps is crucial. We simulated climate warming and top predator loss in natural microcosms to investigate their direct and indirect effects on temporal stability of microbial communities and the role of underlying stabilising mechanisms. Community stability was insensitive to warming, but indirectly decreased due to top predator loss via increased mesopredator abundance and consequent reduction of species asynchrony and species stability. The magnitude of destabilising effects differed among trophic levels, being disproportionally higher at lower trophic levels (e.g. producers). Our study unravels major patterns and causal mechanisms by which trophic downgrading destabilises large food webs, regardless of climate warming scenarios.

Pervasive decline of subtropical aquatic insects over 20 years driven by water transparency, non-native fish and stoichiometric imbalance.

Insect abundance and diversity are declining worldwide. Although recent research found freshwater insect populations to be increasing in some regions, there is a critical lack of data from tropical and subtropical regions. Here, we examine a 20-year monitoring dataset of freshwater insects from a subtropical floodplain comprising a diverse suite of rivers, shallow lakes, channels and backwaters. We found a pervasive decline in abundance of all major insect orders (Odonata, Ephemeroptera, Trichoptera, Megaloptera, Coleoptera, Hemiptera and Diptera) and families, regardless of their functional role or body size. Similarly, Chironomidae species richness decreased over the same time period. The main drivers of this pervasive insect decline were increased concurrent invasions of non-native insectivorous fish, water transparency and changes to water stoichiometry (i.e. N : P ratios) over time. All these drivers represent human impacts caused by reservoir construction. This work sheds light on the importance of long-term studies for a deeper understanding of human-induced impacts on aquatic insects. We highlight that extended anthropogenic impact monitoring and mitigation actions are pivotal in maintaining freshwater ecosystem integrity.

Warming of aquatic ecosystems disrupts aquatic-terrestrial linkages in the tropics.

Aquatic ecosystems are tightly linked to terrestrial ecosystems by exchanges of resources, which influence species interactions, community dynamics and functioning in both ecosystem types. However, our understanding of how this coupling responds to climate warming is restricted to temperate, boreal and arctic regions, with limited knowledge from tropical ecosystems. We investigated how warming aquatic ecosystems impact cross-ecosystem exchanges in the tropics, through the export of aquatic resources into the terrestrial environment and the breakdown of terrestrial resources within the aquatic environment. We experimentally heated 50 naturally assembled aquatic communities, contained within different-sized tank-bromeliads, to a 23.5-32°C gradient of mean water temperatures. The biomass, abundance and richness of aquatic insects emerging into the terrestrial environment all declined with rising temperatures over a 45-day experiment. Structural equation and linear mixed effects modelling suggested that these impacts were driven by deleterious effects of warming on insect development and survival, rather than being mediated by aquatic predation, nutrient availability or reduced body size. Decomposition was primarily driven by microbial activity. However, total decomposition by both microbes and macroinvertebrates increased with temperature in all but the largest ecosystems, where it decreased. Thus, warming decoupled aquatic and terrestrial ecosystems, by reducing the flux of aquatic resources to terrestrial ecosystems but variably enhancing or reducing terrestrial resource breakdown in aquatic ecosystems. In contrast with increased emergence observed in warmed temperate ecosystems, future climate change is likely to reduce connectivity between tropical terrestrial and aquatic habitats, potentially impacting consumers in both ecosystem types. As tropical ectotherms live closer to their thermal tolerance limits compared to temperate species, warming can disrupt cross-ecosystem dynamics in an interconnected tropical landscape and should be considered when investigating ecosystem-level consequences of climate change.

Os ecossistemas aquáticos estão intimamente ligados aos ecossistemas terrestres por meio das trocas de recursos, que influenciam as interações entre as espécies, a dinâmica da comunidade e o funcionamento de ambos os tipos de ecossistemas. No entanto, nosso entendimento de como esse acoplamento responde ao aquecimento do clima é restrito às regiões temperadas, boreais e árticas, com conhecimento limitado para os ecossistemas tropicais. Investigamos como o aquecimento dos ecossistemas aquáticos impacta as trocas entre os ecossistemas nos trópicos, por meio da exportação de recursos aquáticos para o ambiente terrestre e da decomposição dos detritos de origem terrestre no ambiente aquático. Nós aquecemos experimentalmente 50 comunidades aquáticas que habitam tanques de bromélias de diferentes tamanhos, submetidas a um gradiente de temperatura média da água variando de 23,5 a 32°C. Em um experimento de 45 dias, a biomassa, abundância e riqueza de insetos aquáticos emergindo para o ambiente terrestre diminuíram com o aumento da temperatura. Modelos lineares mistos e de equações estruturais sugerem que esses impactos foram causados por efeitos deletérios do aquecimento no desenvolvimento e sobrevivência dos insetos, ao invés de serem mediados por predadores aquáticos, disponibilidade de nutrientes ou tamanho corporal reduzido. A decomposição foi determinada principalmente pela atividade microbiana. A decomposição total por micro-organismos e macro invertebrados aumentou com a temperatura, exceto em ecossistemas maiores. Assim, o aquecimento dissociou os ecossistemas aquáticos e terrestres, reduzindo o fluxo de recursos aquáticos para os ecossistemas terrestres, mas aumentando ou reduzindo de forma variável a decomposição dos recursos terrestres nos ecossistemas aquáticos. Em contraste com o aumento da emergência observada em ecossistemas temperados aquecidos, as mudanças climáticas futuras provavelmente reduzirão a conectividade entre os habitats terrestres e aquáticos tropicais, impactando potencialmente os consumidores em ambos os tipos de ecossistemas. Como organismos ectotérmicos tropicais vivem mais perto dos seus limites de tolerância térmica em comparação com espécies temperadas, o aquecimento pode comprometer a dinâmica entre os ecossistemas em uma paisagem tropical interconectada e deve ser considerado ao investigar as consequências das mudanças climáticas no nível do ecossistema.

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.

Precipitation and predation risk alter the diversity and behavior of pollinators and reduce plant fitness.

Biotic and abiotic factors may individually or interactively disrupt plant-pollinator interactions, influencing plant fitness. Although variations in temperature and precipitation are expected to modify the overall impact of predators on plant-pollinator interactions, few empirical studies have assessed if these weather conditions influence anti-predator behaviors and how this context-dependent response may cascade down to plant fitness. To answer this question, we manipulated predation risk (using artificial spiders) in different years to investigate how natural variation in temperature and precipitation may affect diversity (richness and composition) and behavioral (visitation) responses of flower-visiting insects to predation risk, and how these effects influence plant fitness. Our findings indicate that predation risk and an increase in precipitation independently reduced plant fitness (i.e., seed set) by decreasing flower visitation. Predation risk reduced pollinator visitation and richness, and altered species composition of pollinators. Additionally, an increase in precipitation was associated with lower flower visitation and pollinator richness but did not alter pollinator species composition. However, maximum daily temperature did not affect any component of the pollinator assemblage or plant fitness. Our results indicate that biotic and abiotic drivers have different impacts on pollinator behavior and diversity with consequences for plant fitness components. Even small variation in precipitation conditions promotes complex and substantial cascading effects on plants by affecting both pollinator communities and the outcome of plant-pollinator interactions. Tropical communities are expected to be highly susceptible to climatic changes, and these changes may have drastic consequences for biotic interactions in the tropics.

Ecological response to altered rainfall differs across the Neotropics.

There is growing recognition that ecosystems may be more impacted by infrequent extreme climatic events than by changes in mean climatic conditions. This has led to calls for experiments that explore the sensitivity of ecosystems over broad ranges of climatic parameter space. However, because such response surface experiments have so far been limited in geographic and biological scope, it is not clear if differences between studies reflect geographic location or the ecosystem component considered. In this study, we manipulated rainfall entering tank bromeliads in seven sites across the Neotropics, and characterized the response of the aquatic ecosystem in terms of invertebrate functional composition, biological stocks (total invertebrate biomass, bacterial density) and ecosystem fluxes (decomposition, carbon, nitrogen). Of these response types, invertebrate functional composition was the most sensitive, even though, in some sites, the species pool had a high proportion of drought-tolerant families. Total invertebrate biomass was universally insensitive to rainfall change because of statistical averaging of divergent responses between functional groups. The response of invertebrate functional composition to rain differed between geographical locations because (1) the effect of rainfall on bromeliad hydrology differed between sites, and invertebrates directly experience hydrology not rainfall and (2) the taxonomic composition of some functional groups differed between sites, and families differed in their response to bromeliad hydrology. These findings suggest that it will be difficult to establish thresholds of "safe ecosystem functioning" when ecosystem components differ in their sensitivity to climatic variables, and such thresholds may not be broadly applicable over geographic space. In particular, ecological forecast horizons for climate change may be spatially restricted in systems where habitat properties mediate climatic impacts, and those, like the tropics, with high spatial turnover in species composition.

Different in the dark: The effect of habitat characteristics on community composition and beta diversity in bromeliad microfauna.

The mechanisms which structure communities have been the focus of a large body of research. Here, we address the question if habitat characteristics describing habitat quality may drive changes in community composition and beta diversity of bromeliad-inhabiting microfauna. In our system, changes in canopy cover along an environmental gradient may affect resource availability, disturbance in form of daily water temperature fluctuations and predation, and thus may lead to changes in community structure of bromeliad microfauna through differences in habitat quality along this gradient. Indeed, we observed distinct changes in microfauna community composition along the environmental gradient explained by changes in the extent of daily water temperature fluctuations. We found beta diversity to be higher under low habitat quality (low canopy cover) than under high habitat quality (high canopy cover), which could potentially be explained by a higher relative importance of stochastic processes under low habitat quality. We also partitioned beta diversity into turnover and nestedness components and we found a nested pattern of beta diversity along the environmental gradient, with communities from the lower-quality habitat being nested subsets of communities from the higher-quality habitat. However, this pattern resulted from an increase in microfauna alpha diversity with an increase in habitat quality. By providing insights into microfauna-environment relationships our results contribute to the mechanistic understanding of community dynamics in small freshwater bodies. Here, we highlight the importance of habitat characteristics representing habitat quality in structuring communities, and suggest that this information may help to improve conservation practices of small freshwater ecosystems.

A meta-analysis of predation risk effects on pollinator behaviour.

Flower-visiting animals are constantly under predation risk when foraging and hence might be expected to evolve behavioural adaptations to avoid predators. We reviewed the available published and unpublished data to assess the overall effects of predators on pollinator behaviour and to examine sources of variation in these effects. The results of our meta-analysis showed that predation risk significantly decreased flower visitation rates (by 36%) and time spent on flowers (by 51%) by pollinators. The strength of the predator effects depended neither on predator taxa and foraging mode (sit-and-wait or active hunters) nor on pollinator lifestyle (social vs. solitary). However, predator effects differed among pollinator taxa: predator presence reduced flower visitation rates and time spent on flowers by Squamata, Lepidoptera and Hymenoptera, but not by Diptera. Furthermore, larger pollinators showed weaker responses to predation risk, probably because they are more difficult to capture. Presence of live crab spiders on flowers had weaker effects on pollinator behaviour than presence of dead or artificial crab spiders or other objects (e.g. dead bees, spheres), suggesting that predator crypsis may be effective to some extent. These results add to a growing consensus on the importance of considering both predator and pollinator characteristics from a community perspective.