Metacommunity Scale Research Articles

Evolution of Dispersal in a Predator-Prey Metacommunity

Dispersal is crucial to allowing species inhabiting patchy or spatially subdivided habitats to persist globally despite the possibility of frequent local extinctions. Theoretical studies have repeatedly demonstrated that species that exhibit a regional metapopulation structure and are subject to increasing rates of local patch extinctions should experience strong selective pressures to disperse more rapidly despite the costs such increased dispersal would entail in terms of decreased local fitness. We extend these studies to consider how extinctions arising from predator-prey interactions affect the evolution of dispersal for species inhabiting a metacommunity. Specifically, we investigate how increasing a strong extinction-prone interaction between a predator and prey within local patches affects the evolution of each species' dispersal. We found that for the predator, as expected, evolutionarily stable strategy (ESS) dispersal rates increased monotonically in response to increasing local extinctions induced by strong predator top-down effects. Unexpectedly for the prey, however, ESS dispersal rates displayed a nonmonotonic response to increasing predator-induced extinction rates-actually decreasing for a significant range of values. These counterintuitive results arise from how extinctions resulting from trophic interactions play out at different spatial scales: interactions that increase extinction rates of both species locally can, at the same time, decrease the frequency of interaction between the prey and predator at the metacommunity scale.

Individual‐based trait analyses reveal assembly patterns in tree sapling communities

AbstractQuestionAre trait‐convergence assembly patterns (TCAP) and/or trait‐divergence assembly patterns (TDAP) of tree sapling communities related to microhabitat gradients in forest patches? If so, which traits are related to such patterns? What are the niche mechanisms potentially responsible for community assembly?LocationGrassland–forest mosaic in Serra do Sudeste region, southern Brazil (ca. 30.8°S, 53.1°W).MethodsWe used individual‐based trait information of 1132 tree saplings in 40 plots along gradients of canopy openness and grazing intensity in forest patches in southern Brazil. We used a trait‐based approach, bypassing taxonomic identification, as a first assessment of sapling community assembly in forest patches.ResultsBoth TCAP and TDAP indicate niche mechanisms underlying assembly of tree sapling communities. Specific leaf area (SLA), mean leaf area (LA) and relative abundance of toothed leaf margin (TLM) maximized TCAP. SLA and TLM tended to increase under more closed canopies, whereas LA had higher values at intermediate canopy openness. SLA and LA maximized TDAP and tended to vary more in more closed canopies, whereas SLA and presence of spines, which also maximized TDAP, showed higher variation under lower grazing intensity.ConclusionThe taxon‐free approach was very useful to infer niche mechanisms of tree sapling assembly in forest patches. By adopting an individual‐based trait approach, we assumed that both intra‐specific and intra‐population trait variability were relevant for revealing assembly patterns. We highlight that the use of individual‐based trait data in a metacommunity framework is an excellent way to evaluate TCAP and TDAP at the metacommunity scale, since it takes into account the entire variation of traits throughout communities.

Spatial scale and species identity influence the indigenous–alien diversity relationship in springtails

Although theory underlying the invasion paradox, or the change in the relationship between the richness of alien and indigenous species from negative to positive with increasing spatial scale, is well developed and much empirical work on the subject has been undertaken, most of the latter has concerned plants and to a lesser extent marine invertebrates. Here we therefore examine the extent to which the relationships between indigenous and alien species richness change from the local metacommunity to the interaction neighborhood scales, and the influences of abundance, species identity, and environmental favorability thereon, in springtails, a significant component of the soil fauna. Using a suite of modeling techniques, including generalized least squares and geographically weighted regressions to account for spatial autocorrelation or nonstationarity of the data, we show that the abundance and species richness of both indigenous and alien species at the metacommunity scale respond strongly to declining environmental favorability, represented here by altitude. Consequently, alien and indigenous diversity covary positively at this scale. By contrast, relationships are more complex at the interaction neighborhood scale, with the relationship among alien species richness and/or density and the density of indigenous species varying between habitats, being negative in some, but positive in others. Additional analyses demonstrated a strong influence of species identity, with negative relationships identified at the interaction neighborhood scale involving alien hypogastrurid springtails, a group known from elsewhere to have negative effects on indigenous species in areas where they have been introduced. By contrast, diversity relationships were positive with the other alien species. These results are consistent with both theory and previous empirical findings for other taxa, that interactions among indigenous and alien species change substantially with spatial scale and that environmental favorability may play a key role in explaining the larger scale patterns. However, they also suggest that the interactions may be affected by the identity of the species concerned, especially at the interaction neighborhood scale.

Species dispersal rates alter diversity and ecosystem stability in pond metacommunities

Metacommunity theory suggests that relationships between diversity and ecosystem stability can be determined by the rate of species dispersal among local communities. The predicted relationships, however, may depend upon the relative strength of local environmental processes and disturbance. Here we evaluate the role of dispersal frequency and local predation perturbations in affecting patterns of diversity and stability in pond plankton metacommunities. Pond metacommunities were composed of three mesocosm communities: one of the three communities maintained constant "press" predation from a selective predator, bluegill sunfish (Lepomis macrochirus); the second community maintained "press" conditions without predation; and the third community experienced recurrent "pulsed" predation from bluegill sunfish. The triads of pond communities were connected at either no, low (0.7%/d), or high (20%/d) planktonic dispersal. Richness and composition of zooplankton and stability of plankton biomass and ecosystem productivity were measured at local and regional spatial scales. Dispersal significantly affected diversity such that local and regional biotas at the low dispersal rate maintained the greatest number of species. The unimodal local dispersal-diversity relationship was predator-dependent, however, as selective press predation excluded species regardless of dispersal. Further, there was no effect of dispersal on beta diversity because predation generated local conditions that selected for distinct community assemblages. Spatial and temporal ecosystem stability responded to dispersal frequency but not predation. Low dispersal destabilized the spatial stability of producer biomass but stabilized temporal ecosystem productivity. The results indicate that selective predation can prevent species augmentation from mass effects but has no apparent influence on stability. Dispersal rates, in contrast, can have significant effects on both species diversity and ecosystem stability at multiple spatial scales in metacommunities.

Dispersal scales up the biodiversity–productivity relationship in an experimental source-sink metacommunity

The influence of biodiversity on ecosystem functioning is a major concern of ecological research. However, the biodiversity-ecosystem functioning relationship has very often been studied independently from the mechanisms allowing coexistence. By considering the effects of dispersal and niche partitioning on diversity, the metacommunity perspective predicts a spatial scale-dependence of the shape of the relationship. Here, we present experimental evidence of such scale-dependent patterns. After approximately 500 generations of diversification in a spatially heterogeneous environment, we measured functional diversity (FD) and productivity at both local and regional scales in experimental source-sink metacommunities of the bacterium Pseudomonas fluorescens SBW25. At the regional scale, environmental heterogeneity yielded high levels of FD and we observed a positive correlation between diversity and productivity. At the local scale, intermediate dispersal increased local FD through a mass effect but there was no correlation between diversity and productivity. These experimental results underline the importance of considering the mechanisms maintaining biodiversity and the appropriate spatial scales in understanding its relationship with ecosystem functioning.

Synchrony and Stability of Food Webs in Metacommunities

Synchrony has fundamental but conflicting implications for the persistence and stability of food webs at local and regional scales. In a constant environment, compensatory dynamics between species can maintain food web stability, but factors that synchronize population fluctuations within and between communities are expected to be destabilizing. We studied the dynamics of a food web in a metacommunity to determine how environmental variability and dispersal affect stability by altering compensatory dynamics and average species abundance. When dispersal rate is high, weak correlated environmental fluctuations promote food web stability by reducing the amplitude of compensatory dynamics. However, when dispersal rate is low, weak environmental fluctuations reduce food web stability by inducing intraspecific synchrony across communities. Irrespective of dispersal rate, strong environmental fluctuations disrupt compensatory dynamics and decrease stability by inducing intermittent correlated fluctuations between consumers in local food webs, which reduce both total consumer abundance and predator abundance. Strong correlated environmental fluctuations lead to (i) spatially asynchronous and highly correlated local consumer dynamics when dispersal is low and (ii) spatially synchronous but intermediate local consumer correlation when dispersal is high. By controlling intraspecific synchrony, dispersal mediates the capacity of strong environmental fluctuations to disrupt compensatory dynamics at both local and metacommunity scales.

Stochastic species turnover and stable coexistence in a species-rich, fire-prone plant community.

Understanding the mechanisms that maintain diversity is important for managing ecosystems for species persistence. Here we used a long-term data set to understand mechanisms of coexistence at the local and regional scales in the Cape Floristic Region, a global hotspot of plant diversity. We used a dataset comprising 81 monitoring sites, sampled in 1966 and again in 1996, and containing 422 species for which growth form, regeneration mode, dispersal distance and abundances at both the local (site) and meta-community scales are known. We found that species presence and abundance were stable at the meta-community scale over the 30 year period but highly unstable at the local scale, and were not influenced by species' biological attributes. Moreover, rare species were no more likely to go extinct at the local scale than common species, and that alpha diversity in local communities was strongly influenced by habitat. We conclude that stochastic environmental fluctuations associated with recurrent fire buffer populations from extinction, thereby ensuring stable coexistence at the meta-community scale by creating a “neutral-like” pattern maintained by niche-differentiation.

A metacommunity‐scale comparison of species‐abundance distribution models for plant communities of eastern Australia

Various models of the species‐abundance distribution (SAD) have been proposed to fit empirically derived data however there is no general consensus as to which model provides the best fit. Further, the zero‐sum multinomial SAD model (ZSM) was proposed as a metacommunity model, yet it has not previously been fitted at the metacommunity scale. We note that SAD models based on compound lognormal distributions (such as the Poisson‐lognormal, PLN, and the negative binomial‐lognormal models, NBLN) can also be thought of as metacommunity models, and we compare these with the ZSM when fitted as metacommunity models to SADs of related communities.We collected five datasets in the Sydney Basin, eastern Australia, representing five different types of subtropical/temperate plant communities ranging from closed warm‐temperate rainforest to open wet sclerophyll forest to dry sclerophyll woodland. For each type of plant community, five local communities were identified across the Sydney Basin, and SAD data collected in five randomly placed 0.2‐ha quadrats at each local community. Analysis was performed at two levels: all abundance data from each local community were pooled across each metacommunity and analysed as a single pooled community; and a metacommunity model was fitted to all five local communities of a community type, simultaneously. For the pooled data, we considered the negative‐binomial (NB) and the log‐series (LS) models in addition to ZSM, PLN and NBLN. All five models performed similarly, however the LS had a better fit to three pooled communities and the ZSM and PLN to the remaining two communities. By contrast, the ZSM performed statistically better against the PLN and NBLN when considered as a metacommunity model. We conclude that the ZSM generally provides a more reliable null model for metacommunity abundance data than the lognormal model.

Diversity and dispersal interactively affect predictability of ecosystem function

Theory and small-scale experiments predict that biodiversity losses can decrease the magnitude and stability of ecosystem services such as production and nutrient cycling. Most of this research, however, has been isolated from the immigration and emigration (dispersal) processes that create and maintain diversity in nature. As common anthropogenic drivers of biodiversity change--such as habitat fragmentation, species introductions and climate change--are mediated by these understudied processes, it is unclear how environmental degradation will affect ecosystem services. Here we tested the interactive effects of mobile grazer diversity and dispersal on the magnitude and stability of ecosystem properties in experimental seagrass communities that were either isolated or connected by dispersal corridors. We show that, contrary to theoretical predictions, increasing the number of mobile grazer species in these metacommunities increased the spatial and temporal variability of primary and secondary production. Moreover, allowing grazers to move among and select patches reduced diversity effects on production. Finally, effects of diversity on stability differed qualitatively between patch and metacommunity scales. Our results indicate that declining biodiversity and habitat fragmentation synergistically influence the predictability of ecosystem functioning.

The effects of resource enrichment, dispersal, and predation on local and metacommunity structure

Community structure is the observable outcome of numerous processes. We conducted a laboratory experiment using a microbial model system to disentangle effects of nutrient enrichment, dispersal, and predation on prey species richness and predator abundance at local and metacommunity scales. Prey species included: Chilomonas sp., Colpidium striatum, Colpoda cucullus, Paramecium tetraurelia, P. caudatum, Philodina sp., Spirostomum sp., Tetrahymena thermophila, and Uronema sp., and Stentor coeruleus was the predator used. We hypothesized that: (1) increased basal resources should maintain greater species richness and higher predator abundance; (2) dispersal should maintain greater species richness; and (3) predation should reduce species richness, especially in the high resource treatments relative to no-predator treatments. Our results support all three hypotheses. Further, we show that dispersal affects richness at the local community scale but not at the metacommunity scale. However, predation seems to have major effects at both the local and metacommunity scale. Overall, our results show that effects of resource enrichment, dispersal, and predation were mostly additive rather than interactive, indicating that it may be sometimes easier to understand their effects than generally thought due to complex interactive effects.

METACOMMUNITY INFLUENCES ON COMMUNITY RICHNESS AT MULTIPLE SPATIAL SCALES: A MICROCOSM EXPERIMENT

Large-scale processes are known to be important for patterns of species richness, yet the ways in which local and larger scale processes interact is not clear. I used metacommunities consisting of five interconnected microbial aquatic communities to examine the manner in which processes at different scales affect local and metacommunity richness. Specifically, I manipulated the potential dispersal rate, whether dispersal was localized or global, and variation in initial community composition. A repeated-measures ANOVA showed that a low dispersal rate and intermediate distance dispersal enhanced local richness. Initial assembly variation had no effect on local richness, while a lack of dispersal or global dispersal reduced local richness. At the metacommunity scale, richness was enhanced throughout the time course of the experiment by initial compositional variation and was reduced by high or global dispersal. The effects of dispersal were contingent on the presence of initial compositional variation. The treatments also affected individual species occupancy patterns, with some benefiting from large-scale processes and others being adversely impacted. These results indicate that the effects of dispersal on species richness have a complex relationship with scale and are not solely divisible into "regional" vs. "local" scales. Finally, predictions of the manner in which dispersal rate structures communities appear dependent upon species compositional variation among communities.

Habitat selection determines abundance, richness and species composition of beetles in aquatic communities

Distribution and abundance patterns at the community and metacommunity scale can result from two distinct mechanisms. Random dispersal followed by non-random, site-specific mortality (species sorting) is the dominant paradigm in community ecology, while habitat selection provides an alternative, largely unexplored, mechanism with different demographic consequences. Rather than differential mortality, habitat selection involves redistribution of individuals among habitat patches based on perceived rather than realized fitness, with perceptions driven by past selection. In particular, habitat preferences based on species composition can create distinct patterns of positive and negative covariance among species, generating more complex linkages among communities than with random dispersal models. In our experiments, the mere presence of predatory fishes, in the absence of any mortality, reduced abundance and species richness of aquatic beetles by up to 80% in comparison with the results from fishless controls. Beetle species' shared habitat preferences generated distinct patterns of species richness, species composition and total abundance, matching large-scale field patterns previously ascribed to random dispersal and differential mortality. Our results indicate that landscape-level patterns of distribution and species diversity can be driven to a large extent by habitat selection behaviour, a critical, but largely overlooked, mechanism of community and metacommunity assembly.

Unified spatial scaling of species and their trophic interactions.

Two largely independent bodies of scaling theory address the quantitative relationships between habitat area, species diversity and trophic interactions. Spatial theory within macroecology addresses how species richness scales with area in landscapes, while typically ignoring interspecific interactions. Complexity theory within community ecology addresses how trophic links scale with species richness in food webs, while typically ignoring spatial considerations. Recent studies suggest unifying these theories by demonstrating how spatial patterns influence food-web structure and vice versa. Here, we follow this suggestion by developing and empirically testing a more unified scaling theory. On the basis of power-law species-area relationships, we develop link-area and non-power-law link-species models that accurately predict how trophic links scale with area and species richness of microcosms, lakes and streams from community to metacommunity levels. In contrast to previous models that assume that species richness alone determines the number of trophic links, these models include the species' spatial distribution, and hence extend the domain of complexity theory to metacommunity scales. This generality and predictive success shows how complexity theory and spatial theory can be unified into a much more general theory addressing new domains of ecology.