A rich body of knowledge links biodiversity to ecosystem functioning (BEF), but it is primarily focused on small scales. We review the current theory and identify six expectations for scale ...dependence in the BEF relationship: (1) a nonlinear change in the slope of the BEF relationship with spatial scale; (2) a scale‐dependent relationship between ecosystem stability and spatial extent; (3) coexistence within and among sites will result in a positive BEF relationship at larger scales; (4) temporal autocorrelation in environmental variability affects species turnover and thus the change in BEF slope with scale; (5) connectivity in metacommunities generates nonlinear BEF and stability relationships by affecting population synchrony at local and regional scales; (6) spatial scaling in food web structure and diversity will generate scale dependence in ecosystem functioning. We suggest directions for synthesis that combine approaches in metaecosystem and metacommunity ecology and integrate cross‐scale feedbacks. Tests of this theory may combine remote sensing with a generation of networked experiments that assess effects at multiple scales. We also show how anthropogenic land cover change may alter the scaling of the BEF relationship. New research on the role of scale in BEF will guide policy linking the goals of managing biodiversity and ecosystems.
We address the challenge of scale for biodiversity and ecosystem functioning (BEF) research. We review current theory and identify six expectations for scale dependence in the BEF relationship. We suggest directions for synthesis that combine theoretical and empirical methods and suggest their application to human transformed landscapes.
Abstract The effects of resource subsidies on a recipient ecosystem depends on a variety of factors. Firstly, increasing the supply of limiting nutrients tends to increase the accumulation of biomass ...across different compartments of the recipient food web. Secondly, in the case of aquatic microorganisms, this accumulation may also be contingent on inputs of energy through light, mediating interactions between heterotrophic bacteria and autotrophic algae. Thirdly, macroscopic consumers feed upon microorganisms and recycle nutrients through excretion and egestion, profoundly affecting nutrient cycling. In order to examine the context‐dependency of nutrient cycling in aquatic systems, we performed an experiment in mesocosms where we followed the responses of dissolved nutrients, microorganisms, and larvae of the cosmopolitan mosquito Culex quinquefasciatus (Diptera: Culicidae) to two terrestrial subsidies differing in carbon (C):nutrient, in combination with two light intensities. We found that the interactions amongst bacteria and algae themselves remained generally unimportant after accounting for treatment effects, and that light did mediate some of the effects of nutrient subsidies on microorganisms. We also found that the presence of mosquito larvae was associated with altered concentrations of nitrogen (N) in some treatments, ultimately benefitting algae. Finally, we found that low C:nutrient subsidies generally increased the development rate of mosquito larvae, and that increased light exposure led to higher mosquito survival. Overall, these patterns may be explained by strong nitrogen limitation in our system, causing both group of microorganisms to respond similarly to variations in nutrient supply without direct covariation, and to the key role of N recycling by mosquito larvae in sustaining algal densities in exposed treatments. The context‐dependency in our study may reflect elemental or molecular constraints other than phosphorus. Our experiment thus calls for greater synthesis amongst ecological theories.
Phylogenetic diversity (PD) describes the total amount of phylogenetic distance among species in a community. Although there has been substantial research on the factors that determine community PD, ...exploration of the consequences of PD for ecosystem functioning is just beginning. We argue that PD may be useful in predicting ecosystem functions in a range of communities, from single‐trophic to complex networks. Many traits show a phylogenetic signal, suggesting that PD can estimate the functional trait space of a community, and thus ecosystem functioning. Phylogeny also determines interactions among species, and so could help predict how extinctions cascade through ecological networks and thus impact ecosystem functions. Although the initial evidence available suggests patterns consistent with these predictions, we caution that the utility of PD depends critically on the strength of phylogenetic signals to both traits and interactions. We advocate for a synthetic approach that incorporates a deeper understanding of how traits and interactions are shaped by evolution, and outline key areas for future research. If these complexities can be incorporated into future studies, relationships between PD and ecosystem function bear promise in conceptually unifying evolutionary biology with ecosystem ecology.
Predators and prey interact at small spatial scales, but during their lifetime disperse at much larger spatial scales. Trophic metacommunity theory proposes that dispersal is a critical process that ...determines food web structure at small and large scales. The application of metacommunity theory to empirical systems remains elusive because key parameters such as dispersal and interaction strengths have been very difficult to quantify. Here we develop a novel approach that combines population genomics with mesocosm experiments to parameterize a metacommunity model. Using genotyping-by-sequencing, we characterized the dispersal kernels of a predator–prey pair living in a phytotelm metacommunity. We found that the prey dispersed up to 25 km while the predator dispersed only 350 m. We then quantified a functional response for these species using feeding trials. Even without invoking differences in the abiotic niche, our empirically parameterized simulation model produced patterns of population survival and occupancy that were consistent with past observations on the natural system. Importantly we found that these patterns were more likely to be found with simulations based on our observed values than in other regions of potential parameter space. This suggests that the observed dispersal kernels contribute to the dynamics of these species in the metacommunity.
Species and trophic richness often increase with habitat size. Although many ecological processes have been evoked to explain both patterns, the environmental stress associated with small habitats ...has rarely been considered. We propose that larger habitats may be species rich simply because their environmental conditions are within the fundamental niche of more species; larger habitats may also have more trophic levels if traits of predators render them vulnerable to environmental stress. We test this hypothesis using the aquatic insect larvae in water-filled bromeliads. In bromeliads, the probability of desiccation is greatest in small plants. For the 10 most common bromeliad insect taxa, we ask whether differences in drought tolerance and regional abundances between taxa predict community and trophic composition over a gradient of bromeliad size. First, we used bromeliad survey data to calculate the mean habitat size of occurrence of each taxon. Comparing the observed mean habitat size of occurrence to that expected from random species assembly based on differences in their regional abundances allowed us to obtain habitat size sensitivity indices (as
Z
scores) for the various insect taxa. Second, we obtained drought sensitivity indices by subjecting individual insects to drought and measuring the effects on relative growth rates in a mesocosm experiment. We found that drought sensitivity strongly predicts habitat size sensitivity in bromeliad insects. However, an increase in trophic richness with habitat size could not be explained by an increased sensitivity of predators to drought, but rather by sampling effects, as predators were rare compared to lower trophic levels. This finding suggests that physiological tolerance to environmental stress can be relevant in explaining the universal increase in species with habitat size.
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.
Accelerating rates of species extinction have prompted a growing number of researchers to manipulate the richness of various groups of organisms and examine how this aspect of diversity impacts ...ecological processes that control the functioning of ecosystems. We summarize the results of 44 experiments that have manipulated the richness of plants to examine how plant diversity affects the production of biomass. We show that mixtures of species produce an average of 1.7 times more biomass than species monocultures and are more productive than the average monoculture in 79% of all experiments. However, in only 12% of all experiments do diverse polycultures achieve greater biomass than their single most productive species. Previously, a positive net effect of diversity that is no greater than the most productive species has been interpreted as evidence for selection effects, which occur when diversity maximizes the chance that highly productive species will be included in and ultimately dominate the biomass of polycultures. Contrary to this, we show that although productive species do indeed contribute to diversity effects, these contributions are equaled or exceeded by species complementarity, where biomass is augmented by biological processes that involve multiple species. Importantly, both the net effect of diversity and the probability of polycultures being more productive than their most productive species increases through time, because the magnitude of complementarity increases as experiments are run longer. Our results suggest that experiments to date have, if anything, underestimated the impacts of species extinction on the productivity of ecosystems.
A central current debate in community ecology concerns the relative importance of deterministic versus stochastic processes underlying community structure. However, the concept of stochasticity ...presents several profound philosophical, theoretical and empirical challenges, which we address here. The philosophical argument that nothing in nature is truly stochastic can be met with the following operational concept of neutral stochasticity in community ecology: change in the composition of a community (i.e. community dynamics) is neutrally stochastic to the degree that individual demographic events – birth, death, immigration, emigration – which cause such changes occur at random with respect to species identities. Empirical methods for identifying the stochastic component of community dynamics or structure include null models and multivariate statistics on observational species-by-site data (with or without environmental or trait data), and experimental manipulations of 'stochastic' species colonization order or relative densities and frequencies of competing species. We identify the fundamental limitations of each method with respect to its ability to allow inferences about stochastic community processes. Critical future needs include greater precision in articulating the link between results and ecological inferences, a comprehensive theoretical assessment of the interpretation of statistical analyses of observational data, and experiments focusing on community size and on natural variation in species colonization order.
Predators and prey often differ in body mass. The ratio of predator to prey body mass influences the predator's functional response (how consumption varies with prey density), and therefore, the ...strength and stability of the predator-prey interaction. The persistence of food chains is maximized when prey species are neither too big nor too small relative to their predator. Nonetheless, we do not know if (i) food web persistence requires that all predator-prey body mass ratios are intermediate, nor (ii) if this constraint depends on prey diversity. We experimentally quantified the functional response for a single predator consuming prey species of different body masses. We used the resultant allometric functional response to parametrize a food web model. We found that predator persistence was maximized when the minimum prey size in the community was intermediate, but as prey diversity increased, the minimum body size could take a broader range of values. This last result occurs because of Jensen's inequality: the average handling time for multiple prey of different sizes is higher than the handling time of the average sized prey. Our results demonstrate that prey diversity mediates how differences between predators and prey in body mass determine food web stability.
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 facultative 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.
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