Nicholas A. C. Marino

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.

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.

Aquatic macroinvertebrate community composition in tank-bromeliads is determined by bromeliad species and its constrained characteristics

Insect herbivore‐plant associations have been widely explored in the scientific literature over the past decades, but other insect‐plant associations have received little focus so far. Tank‐bromeliads host a large number of invertebrate species and, even though previous studies showed a strong variation in community composition among tank‐bromeliad species, the reasons for this variation are still unclear. In this study, we first examined whether different bromeliad species supports distinct and predictable community compositions and whether such variation was caused by (i) the bromeliad species itself, (ii) to its environment, and/or (iii) the covariance between both. We found that different tank‐bromeliad species hosts distinct macroinvertebrate assemblages, but community composition was particularly less predictable in the smaller tank‐bromeliads. Aquatic macroinvertebrates were sorted along a gradient of environmental conditions presented by the set of tank‐bromeliad species examined, according to their functional and biological traits. The tank‐bromeliad species and the environmental conditions within its tanks were strongly related and equally important in structuring these communities. These results suggest that associations between bromeliads and the fauna inhabiting its tanks are likely a result of the environmental characteristics particular to each bromeliad species.

Habitat size determine algae biomass in tank-bromeliads

The primary goal of this study was to evaluate the relative importance of habitat physical properties, bottom-up and top-down factors, and their interaction on algae biomass in tank-bromeliads. We sampled algae biomass (chlorophyll-a concentration), micro-metazoan density, mosquito abundance, and several environmental variables, including nutrient concentrations and characteristics of the habitat physical structure, in a survey of 64 tank-bromeliads of four different species (Aechmea nudicaulis, Aechmea lingulata, Neoregelia cruenta, and Vriesea neoglutinosa). We analyzed the complete and individual bromeliad species datasets using an information-theoretic model selection approach (Akaike’s information criterion). Bromeliad species, maximum water volume, and bromeliad diameter comprised the best model for determining chlorophyll-a concentrations for the complete dataset. The maximum water volume also comprised the best model to explain chlorophyll-a concentrations in three of four bromeliad species datasets. Interactions between consumers and nutrient concentration were included in the subsequent models, but they were not statistically significant. Taken together, these results demonstrate that the impact of habitat size on the associated autotrophic biomass occurs possibly via changes in community susceptibility to disturbances, particularly drought. We can conclude that habitat size is more important than resource availability or herbivory on phytotelm autotrophic biomass regulation in these natural microcosms.

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.

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.

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.

Environmental control of the microfaunal community structure in tropical bromeliads

Abstract Ecological communities hosted within phytotelmata (plant compartments filled with water) provide an excellent opportunity to test ecological theory and to advance our understanding of how local and global environmental changes affect ecosystems. However, insights from bromeliad phytotelmata communities are currently limited by scarce accounts of microfauna assemblages, even though these assemblages are critical in transferring, recycling, and releasing nutrients in these model ecosystems. Here, we analyzed natural microfaunal communities in leaf compartments of 43 bromeliads to identify the key environmental filters underlying their community structures. We found that microfaunal community richness and abundance were negatively related to canopy openness and vertical height above the ground. These associations were primarily driven by the composition of amoebae and flagellate assemblages and indicate the importance of bottom‐up control of microfauna in bromeliads. Taxonomic richness of all functional groups followed a unimodal relationship with water temperature, peaking at 23–25°C and declining below and above this relatively narrow thermal range. This suggests that relatively small changes in water temperature under expected future climate warming may alter taxonomic richness and ecological structure of these communities. Our findings improve the understanding of this unstudied but crucial component of bromeliad ecosystems and reveal important environmental filters that likely contribute to overall bromeliad community structure and function.

Geographical and experimental contexts modulate the effect of warming on top‐down control: a meta‐analysis

Ecologists have extensively investigated the effect of warming on consumer-resource interactions, with experiments revealing that warming can strengthen, weaken or have no net effect on top-down control of resources. These experiments have inspired a body of theoretical work to explain the variation in the effect of warming on top-down control. However, there has been no quantitative attempt to reconcile theory with outcomes from empirical studies. To address the gap between theory and experiment, we performed a meta-analysis to examine the combined effect of experimental warming and top-down control on resource biomass and determined potential sources of variation across experiments. We show that differences in experimental outcomes are related to systematic variation in the geographical distribution of studies. Specifically, warming strengthened top-down control when experiments were conducted in colder regions, but had the opposite effect in warmer regions. Furthermore, we found that differences in the thermoregulation strategy of the consumer and openness of experimental arenas to dispersal can contribute to some deviation from the overall geographical pattern. These results reconcile empirical findings and support the expectation of geographical variation in the response of consumer-resource interactions to warming.