Species and individual replacements contribute more than nestedness to shape vertebrate scavenger metacommunities

Understanding the mechanisms that organize biodiversity is central in ecology and conservation. Beta diversity links local (alfa) and regional (gamma) diversity, giving insight into how communities organize spatially. Metacommunity ecology provides the framework to interpret regional and local processes interacting to shape communities. However, the lack of metacommunity studies for large vertebrates may limit the understanding and compromise the preservation of ecosystem functions and services. We aim to understand the mechanisms underlying differences in species composition among vertebrate scavenger communities ? which provide key ecosystem functions, e.g. carrion consumption ? within a metacommunity context. We obtained species richness and abundances at scavenger communities consuming ungulate carcasses monitored through motion‐triggered remote cameras in seven terrestrial ecosystems in Spain. We partitioned beta diversity to decompose incidence‐based (species presence/absence) and abundance‐based dissimilarities into their components (turnover/balanced variation and nestedness/abundance gradient, respectively). We identified the environmental factors explaining the observed patterns. The vertebrate scavenger metacommunity consisted of 3101 individuals from 30 species. Changes in composition among ecosystems were mostly (> 84%) due to species or individual replacement (i.e. turnover or balanced variation). Species or individual loss/gain (i.e. nestedness or abundance gradient) accounted for 13–16% of these changes. Mean carcass weight, elevation and habitat diversity were the main factors explaining species/individual replacement. Our findings suggest that local processes such as species‐sorting through habitat heterogeneity would dominate scavenger metacommunity dynamics together with stochastic forces (i.e. related to carrion unpredictability and scavenging being a widespread strategy among vertebrates). The presence of structured patterns (i.e. nestedness) in beta diversity could reflect a role of deterministic processes: mass‐effects through dispersal and defaunation. Vultures are long‐distance foragers and functionally dominant species, which would connect local assemblages within the metacommunity, supporting scavenger diversity and functions across space. These results highlight the importance of managing vertebrate scavenger assemblages within a metacommunity context.     

Using functional diversity patterns to explore metacommunity dynamics: a framework for understanding local and regional influences on community structure

The metacommunity concept, describing how local and regional scale processes interact to structure communities, has been successfully applied to patterns of taxonomic diversity. Functional diversity has proved useful for understanding local scale processes, but has less often been applied to understanding regional scale processes. Here, we explore functional diversity patterns within a metacommunity context to help elucidate how local and regional scale processes influence community assembly. We detail how each of the four metacommunity perspectives (species sorting, mass effects, patch dynamics, neutral) predict different patterns of functional beta‐ and alpha‐diversity and spatial structure along two key gradients: dispersal limitation and environmental conditions. We then apply this conceptual model to a case study from alpine tundra plant communities. We sampled species composition in 17 ‘sky islands’ of alpine tundra in the Colorado Rocky Mountains, USA that differed in geographic isolation and area (key factors related to dispersal limitation) and temperature and elevation (key environmental factors). We quantified functional diversity in each site based on specific leaf area, leaf area, stomatal conductance, plant height and chlorophyll content. We found that colder high elevation sites were functionally more similar to each other (decreased functional beta‐diversity) and had lower functional alpha‐diversity. Geographic isolation and area did not influence functional beta‐ or alpha‐diversity. These results suggest a strong role for environmental conditions structuring alpine plant communities, patterns consistent with the species sorting metacommunity perspective. Incorporating functional diversity into metacommunity theory can help elucidate how local and regional factors structure communities and provide a framework for observationally examining the role of metacommunity dynamics in systems where experimental approaches are less tractable.     

A comparative analysis of metacommunity types in the freshwater realm

Most metacommunity studies have taken a direct mechanistic approach, aiming to model the effects of local and regional processes on local communities within a metacommunity. An alternative approach is to focus on emergent patterns at the metacommunity level through applying the elements of metacommunity structure (EMS; Oikos, 97, 2002, 237) analysis. The EMS approach has very rarely been applied in the context of a comparative analysis of metacommunity types of main microbial, plant, and animal groups. Furthermore, to our knowledge, no study has associated metacommunity types with their potential ecological correlates in the freshwater realm. We assembled data for 45 freshwater metacommunities, incorporating biologically highly disparate organismal groups (i.e., bacteria, algae, macrophytes, invertebrates, and fish). We first examined ecological correlates (e.g., matrix properties, beta diversity, and average characteristics of a metacommunity, including body size, trophic group, ecosystem type, life form, and dispersal mode) of the three elements of metacommunity structure (i.e., coherence, turnover, and boundary clumping). Second, based on those three elements, we determined which metacommunity types prevailed in freshwater systems and which ecological correlates best discriminated among the observed metacommunity types. We found that the three elements of metacommunity structure were not strongly related to the ecological correlates, except that turnover was positively related to beta diversity. We observed six metacommunity types. The most common were Clementsian and quasi‐nested metacommunity types, whereas Random, quasi‐Clementsian, Gleasonian, and quasi‐Gleasonian types were less common. These six metacommunity types were best discriminated by beta diversity and the first axis of metacommunity ecological traits, ranging from metacommunities of producer organisms occurring in streams to those of large predatory organisms occurring in lakes. Our results showed that focusing on the emergent properties of multiple metacommunities provides information additional to that obtained in studies examining variation in local community structure within a metacommunity.     

Evaluating the role of regional and local processes in structuring a larval trematode metacommunity of Helisoma trivolvis

Metacommunity theory has advanced our understanding of how local and regional processes affect the structure of ecological communities. While parasites have largely been omitted from metacommunity research, parasite communities can provide the large sample sizes and discrete boundaries often required for evaluating metacommunity patterns. Here, we used assemblages of flatworm parasites that infect freshwater snails (Helisoma trivolvis) to evaluate three questions: 1) what factors affect individual host infections within ponds? 2) Is the parasite metacommunity structured among ponds? And 3) what is the relative role of local versus regional processes in determining metacommunity structure and species richness among ponds? We examined 10 821 snails from 96 sites in five park complexes in the San Francisco Bay area, California, and found 953 infections from six parasite groups. At the within‐pond level, infection status of host snails correlated positively with individual snail size and pond infection prevalence for all six parasite groups. Using an ordination method to test for metacommunity structure, we found that the parasite metacommunity was organized in a non‐random pattern with species responding individually along an environmental gradient. Based on a model selection approach involving local and regional predictors, parasite species richness and metacommunity structure correlated with both local abiotic (pH and total dissolved nitrogen) and biotic (non‐host mollusk density, and H. trivolvis biomass) factors, with little support for regional predictors. Overall, this trematode metacommunity most closely followed the predictions from the species sorting or mass effects metacommunity paradigm, in which community diversity is filtered by local site characteristics.     

Contribution of neutral processes to the assembly of gut microbial communities in the zebrafish over host development

Despite their importance to host health and development, the communities of microorganisms associated with humans and other animals are characterized by a large degree of unexplained variation across individual hosts. The processes that drive such inter-individual variation are not well understood. To address this, we surveyed the microbial communities associated with the intestine of the zebrafish, Danio rerio, over developmental time. We compared our observations of community composition and distribution across hosts with that predicted by a neutral assembly model, which assumes that community assembly is driven solely by chance and dispersal. We found that as hosts develop from larvae to adults, the fit of the model to observed microbial distributions decreases, suggesting that the relative importance of non-neutral processes, such as microbe-microbe interactions, active dispersal, or selection by the host, increases as hosts mature. We also observed that taxa which depart in their distributions from the neutral prediction form ecologically distinct sub-groups, which are phylogenetically clustered with respect to the full metacommunity. These results demonstrate that neutral processes are sufficient to generate substantial variation in microbiota composition across individual hosts, and suggest that potentially unique or important taxa may be identified by their divergence from neutral distributions.     

A meta‐analysis suggesting that the relationship between biodiversity and risk of zoonotic pathogen transmission is idiosyncratic

Zoonotic pathogens are significant burdens on global public health. Because they are transmitted to humans from non‐human animals, the transmission dynamics of zoonoses are necessarily influenced by the ecology of their animal hosts and vectors. The ‘dilution effect’ proposes that increased species diversity reduces disease risk, suggesting that conservation and public health initiatives can work synergistically to improve human health and wildlife biodiversity. However, the meta‐analysis that we present here indicates a weak and highly heterogeneous relationship between host biodiversity and disease. Our results suggest that disease risk is more likely a local phenomenon that relies on the specific composition of reservoir hosts and vectors, and their ecology, rather than patterns of species biodiversity.     

Ecosystem dynamics, biological diversity and emerging infectious diseases

In this article, we summarize the major scientific developments of the last decade on the transmission of infectious agents in multi-host systems. Almost sixty percent of the pathogens that have emerged in humans during the last 30-40 years are of animal origin and about sixty percent of them show an important variety of host species besides humans (3 or more possible host species). In this review, we focus on zoonotic infections with vector-borne transmission and dissect the contrasting effects that a multiplicity of host reservoirs and vectors can have on their disease dynamics. We discuss the effects exerted by host and vector species richness and composition on pathogen prevalence (i.e., reduction, including the dilution effect, or amplification). We emphasize that, in multiple host systems and for vector-borne zoonotic pathogens, host reservoir species and vector species can exert contrasting effect locally. The outcome on disease dynamics (reduced pathogen prevalence in vectors when the host reservoir species is rich and increased pathogen prevalence when the vector species richness increases) may be highly heterogeneous in both space and time. We then ask briefly how a shift towards a more systemic perspective in the study of emerging infectious diseases, which are driven by a multiplicity of hosts, may stimulate further research developments. Finally, we propose some research avenues that take better into account the multi-host species reality in the transmission of the most important emerging infectious diseases, and, particularly, suggest, as a possible orientation, the careful assessment of the life-history characteristics of hosts and vectors in a community ecology-based perspective.     

Inferring species roles in metacommunity structure from species co-occurrence networks

A long-standing question in community ecology is what determines the identity of species that coexist across local communities or metacommunity assembly. To shed light upon this question, we used a network approach to analyse the drivers of species co-occurrence patterns. In particular, we focus on the potential roles of body size and trophic status as determinants of metacommunity cohesion because of their link to resource use and dispersal ability. Small-sized individuals at low-trophic levels, and with limited dispersal potential, are expected to form highly linked subgroups, whereas large-size individuals at higher trophic positions, and with good dispersal potential, will foster the spatial coupling of subgroups and the cohesion of the whole metacommunity. By using modularity analysis, we identified six modules of species with similar responses to ecological conditions and high co-occurrence across local communities. Most species either co-occur with species from a single module or are connectors of the whole network. Among the latter are carnivorous species of intermediate body size, which by virtue of their high incidence provide connectivity to otherwise isolated communities playing the role of spatial couplers. Our study also demonstrates that the incorporation of network tools to the analysis of metacommunity ecology can help unveil the mechanisms underlying patterns and processes in metacommunity assembly.     

No evidence for environmental and spatial processes in structuring phytoplankton communities

The relative importance of local and regional processes in shaping natural communities within a metacommunity context has been a focus of intense debate in recent years. Floodplain lakes provide a good system for testing this theoretical approach, as they undergo seasonal variations in physical, chemical and biological factors, as well as in their degree of connectivity. Here, we investigated how local phytoplankton communities in lakes of a tropical river-floodplain system (Araguaia River floodplain – Central Brazil) were affected by environmental and spatial (dispersal) predictors in two rainy and two dry seasons (two consecutive years). Partial redundancy analysis indicated that during the periods analyzed the effects of neither predictor were significant. Although we cannot exclude the possibility that these tropical phytoplankton communities could be regulated by stochastic events, we suggested that further studies will have greater explanatory power if they include other variables related to biotic interactions (e.g., abundance of grazers) and fine-scale environmental variation.     

Local factors determine plant community structure on closely neighbored islands

Despite the recent popularity of the metacommunity concept, ecologists have not evaluated the applicability of different metacommunity frameworks to insular organisms. We surveyed 50 closely spaced islands in the Thousand-Island Lake of China to examine the role of local (environmental) and regional (dispersal) factors in structuring woody plant assemblages (tree and shrub species) on these islands. By partitioning the variation in plant community structure into local and regional causes, we showed that local environmental conditions, specifically island morphometric characteristics, accounted for the majority of the variation in plant community structure among the studied islands. Spatial variables, representing the potential importance of species dispersal, explained little variation. We conclude that one metacommunity framework-species sorting-best characterizes these plant communities. This result reinforces the idea that the traditional approach of emphasizing the local perspective when studying ecological communities continues to hold its value.     

Fine-scale variation in vector host use and force of infection drive localized patterns of West Nile virus transmission

The influence of host diversity on multi-host pathogen transmission and persistence can be confounded by the large number of species and biological interactions that can characterize many transmission systems. For vector-borne pathogens, the composition of host communities has been hypothesized to affect transmission; however, the specific characteristics of host communities that affect transmission remain largely unknown. We tested the hypothesis that vector host use and force of infection (i.e., the summed number of infectious mosquitoes resulting from feeding upon each vertebrate host within a community of hosts), and not simply host diversity or richness, determine local infection rates of West Nile virus (WNV) in mosquito vectors. In suburban Chicago, Illinois, USA, we estimated community force of infection for West Nile virus using data on Culex pipiens mosquito host selection and WNV vertebrate reservoir competence for each host species in multiple residential and semi-natural study sites. We found host community force of infection interacted with avian diversity to influence WNV infection in Culex mosquitoes across the study area. Two avian species, the American robin (Turdus migratorius) and the house sparrow (Passer domesticus), produced 95.8% of the infectious Cx. pipiens mosquitoes and showed a significant positive association with WNV infection in Culex spp. mosquitoes. Therefore, indices of community structure, such as species diversity or richness, may not be reliable indicators of transmission risk at fine spatial scales in vector-borne disease systems. Rather, robust assessment of local transmission risk should incorporate heterogeneity in vector host feeding and variation in vertebrate reservoir competence at the spatial scale of vector-host interaction.     

Spatiotemporal dynamics in a seasonal metacommunity structure is predictable: the case of floodplain‐fish communities

The metacommunity framework has greatly advanced our understanding about the importance of local and regional processes structuring ecological communities. However, information on how metacommunity structure and the relative strengths of their underlying mechanisms change through time is largely lacking. Dynamic systems that undergo environmental temporal changes and disturbances, such as floodplains, serve as natural laboratories to explore how their metacommunity structure change in time. Here we applied the Elements of Metacommunity Structure framework and variation partitioning analysis to assess how temporal changes in the local environmental factors and regional dispersal processes in the rain season influence a seasonal floodplain‐fish metacommunity. Across four months, relevant environmental factors were measured across 21 patches where over 3500 individual fish were sampled. Connectivity was measured using landscape resistance‐based metrics and additional spatial variation in metacommunity structure was assessed via spatial autocorrelation functions. The metacommunity structure changed from nestedness, at the beginning of the flood season, to a quasi‐Clementsian gradient at the end. Our analyses show that connectivity is only important in the beginning of the flood season whereas environment is only important at the end. These results suggest that this metacommunity is structured by changes between dispersal limitation and environmental filtering through time.     

Diseases of humans and their domestic mammals: pathogen characteristics, host range and the risk of emergence

Pathogens that can be transmitted between different host species are of fundamental interest and importance from public health, conservation and economic perspectives, yet systematic quantification of these pathogens is lacking. Here, pathogen characteristics, host range and risk factors determining disease emergence were analysed by constructing a database of disease-causing pathogens of humans and domestic mammals. The database consisted of 1415 pathogens causing disease in humans, 616 in livestock and 374 in domestic carnivores. Multihost pathogens were very prevalent among human pathogens (61.6%) and even more so among domestic mammal pathogens (livestock 77.3%, carnivores 90.0%). Pathogens able to infect human, domestic and wildlife hosts contained a similar proportion of disease-causing pathogens for all three host groups. One hundred and ninety-six pathogens were associated with emerging diseases, 175 in humans, 29 in livestock and 12 in domestic carnivores. Across all these groups, helminths and fungi were relatively unlikely to emerge whereas viruses, particularly RNA viruses, were highly likely to emerge. The ability of a pathogen to infect multiple hosts, particularly hosts in other taxonomic orders or wildlife, were also risk factors for emergence in human and livestock pathogens. There is clearly a need to understand the dynamics of infectious diseases in complex multihost communities in order to mitigate disease threats to public health, livestock economies and wildlife.     

The Relationship between Host Lifespan and Pathogen Reservoir Potential: An Analysis in the System Arabidopsis thaliana-Cucumber mosaic virus

Identification of the determinants of pathogen reservoir potential is central to understand disease emergence. It has been proposed that host lifespan is one such determinant: short-lived hosts will invest less in costly defenses against pathogens, so that they will be more susceptible to infection, more competent as sources of infection and/or will sustain larger vector populations, thus being effective reservoirs for the infection of long-lived hosts. This hypothesis is sustained by analyses of different hosts of multihost pathogens, but not of different genotypes of the same host species. Here we examined this hypothesis by comparing two genotypes of the plant Arabidopsis thaliana that differ largely both in life-span and in tolerance to its natural pathogen Cucumber mosaic virus (CMV). Experiments with the aphid vector Myzus persicae showed that both genotypes were similarly competent as sources for virus transmission, but the short-lived genotype was more susceptible to infection and was able to sustain larger vector populations. To explore how differences in defense against CMV and its vector relate to reservoir potential, we developed a model that was run for a set of experimentally-determined parameters, and for a realistic range of host plant and vector population densities. Model simulations showed that the less efficient defenses of the short-lived genotype resulted in higher reservoir potential, which in heterogeneous host populations may be balanced by the longer infectious period of the long-lived genotype. This balance was modulated by the demography of both host and vector populations, and by the genetic composition of the host population. Thus, within-species genetic diversity for lifespan and defenses against pathogens will result in polymorphisms for pathogen reservoir potential, which will condition within-population infection dynamics. These results are relevant for a better understanding of host-pathogen co-evolution, and of the dynamics of pathogen emergence.     

A process‐based metacommunity framework linking local and regional scale community ecology

The metacommunity concept has the potential to integrate local and regional dynamics within a general community ecology framework. To this end, the concept must move beyond the discrete archetypes that have largely defined it (e.g. neutral vs. species sorting) and better incorporate local scale species interactions and coexistence mechanisms. Here, we present a fundamental reconception of the framework that explicitly links local coexistence theory to the spatial processes inherent to metacommunity theory, allowing for a continuous range of competitive community dynamics. These dynamics emerge from the three underlying processes that shape ecological communities: (1) density‐independent responses to abiotic conditions, (2) density‐dependent biotic interactions and (3) dispersal. Stochasticity is incorporated in the demographic realisation of each of these processes. We formalise this framework using a simulation model that explores a wide range of competitive metacommunity dynamics by varying the strength of the underlying processes. Using this model and framework, we show how existing theories, including the traditional metacommunity archetypes, are linked by this common set of processes. We then use the model to generate new hypotheses about how the three processes combine to interactively shape diversity, functioning and stability within metacommunities.     

Linking metacommunity theory and symbiont evolutionary ecology

Processes that occur both within and between hosts can influence the ecological and evolutionary dynamics of symbionts, a broad term that includes parasitic and disease-causing organisms. Metacommunity theory can integrate these local- and regional-scale dynamics to explore symbiont community composition patterns across space. In this article I emphasize that symbionts should be incorporated into the metacommunity concept. I highlight the utility of metacommunity theory by discussing practical and general benefits that emerge from considering symbionts in a metacommunity framework. Specifically, investigating the local and regional drivers of symbiont community and metacommunity structure will lead to a more holistic understanding of symbiont ecology and evolution and could reveal novel insights into the roles of symbiont communities in mediating host health.     

Effects of a zoonotic pathogen,Borrelia burgdorferi,on the behavior of a key reservoir host

Most emerging infectious diseases of humans are transmitted to humans from other animals. The transmission of these “zoonotic” pathogens is affected by the abundance and behavior of their wildlife hosts. However, the effects of infection with zoonotic pathogens on behavior of wildlife hosts, particularly those that might propagate through ecological communities, are not well understood. Borrelia burgdorferi is a bacterium that causes Lyme disease, the most common vector‐borne disease in the USA and Europe. In its North American range, the pathogen is most frequently transmitted among hosts through the bite of infected blacklegged ticks (Ixodes scapularis). Using sham and true vaccines, we experimentally manipulated infection load with this zoonotic pathogen in its most competent wildlife reservoir host, the white‐footed mouse, Peromyscus leucopus, and quantified the effects of infection on mouse foraging behavior, as well as levels of mouse infestation with ticks. Mice treated with the true vaccine had 20% fewer larval blacklegged ticks infesting them compared to mice treated with the sham vaccine, a significant difference. We observed a nonsignificant trend for mice treated with the true vaccine to be more likely to visit experimental foraging trays (20%–30% effect size) and to prey on gypsy moth pupae (5%–20% effect size) compared to mice treated with the sham vaccine. We observed no difference between mice on true‐ versus sham‐vaccinated grids in risk‐averse foraging. Infection with this zoonotic pathogen appears to elicit behavioral changes that might reduce self‐grooming, but other behaviors were affected subtly or not at all. High titers of B. burgdorferi in mice could elicit a self‐reinforcing feedback loop in which reduced grooming increases tick burdens and hence exposure to tick‐borne pathogens.     

Assessing the effect of zooprophylaxis on zoonotic cutaneous leishmaniasis transmission: A system dynamics approach

Capturing or diverting the disease carrying vector from humans can reduce the transmission of vector borne diseases such as leishmaniasis. The use of animals that act as dead-end hosts to relieve the vector (sandfly) bites on humans is called zooprophylaxis. However, as the number of blood meal providers especially domestic animals increases, the sandflies enhanced availability of blood meals will improve its number and survival, thereby countering the impact of diverting bites from humans. Thus, the transmission model exhibits the structure of a feedback loop characterizing complex dynamic systems. In order to rigorously assess the effect of zooprophylaxis, we propose a system dynamic model for zoonotic cutaneous leishmaniasis transmission with 3 blood-meal hosts: domestic animals, humans, and a reservoir (rodents). In this context, a simulation study of the proposed model with a follow-up period of 1000 days was performed.     

Disentangling the role of connectivity,environmental filtering,and spatial structure on metacommunity dynamics

Dispersal is a key process in metacommunity dynamics, allowing the maintenance of diversity in complex community networks. Geographic distance is usually used as a surrogate for connectivity implying that communities that are closely located are considered more prone to exchange individuals than distant communities. However, in some natural systems, organisms may be subjected to directional dispersal (air or water flows, particular landscape configuration), possibly leading close communities to be isolated from each other and distant communities to be connected. Using geographic distance as a proxy for realised connectivity may then yield misleading results regarding the role of dispersal in structuring communities in such systems. Here, we quantified the relative importance of flow connectivity, geographic distance, and environmental gradients to explain polychaete metacommunity structure along the coasts of the Gulf of Lions (northwest Mediterranean Sea). Flow connectivity was estimated by Lagrangian particle dispersal simulations. Our results revealed that this metacommunity is strongly structured by the environment at large spatial scales, and that both flow connectivity and geographic distance play an important role within homogeneous environments at smaller spatial scales. We thus strongly advocate for a wider use of connectivity measures, in addition to geographic distance, to study spatial patterns of biological diversity (e.g. distance decay) and to infer the processes behind these patterns at different spatial scales. Synthesis Everything is connected, but connections are seldom accurately quantified. Biological communities are often studied separately, using observations, experiments and models to unravel local dynamics of organisms interacting with each other. However, regional processes such as dispersal through ocean and air circulation, likely to connect distant communities and influence their local dynamics, are not always accounted for, or, at best, used as an homogeneous and distance‐related factor. Ocean models have being extensively developed and validated during the past decades with the increasing availability of accurate meteorological data. Using such model outputs, precise quantifi cation of exchange rates of organisms between communities was performed in a marine Mediterranean coastal area. Jointly with local environmental and biological data, these results were used to quantify the effects of realistic connectivity on local and regional polychaete community structure, and revealed that the environmental gradient, geographic distance, and connectivity were responsible for community structure at different spatial scales.     

The influence of dispersal on the realized trajectory of a pond metacommunity

Dispersal rates play a critical role in metacommunity dynamics, yet few studies have attempted to characterize dispersal rates for the majority of species in any natural community. Here we evaluate the relationship between the abundances of 179 plankton taxa in a pond metacommunity and their dispersal rates. We find the expected positive relationship between the regional abundances of phytoplankton, protozoa and metazoan zooplankton, which is suggestive of dispersal being a density‐independent per capita rate for these groups. When we tested to see if the rates of dispersing taxa predicted changes in community composition, we found that dispersers had no measurable impact on the short‐term trajectory of local pond communities or mesocosm communities established experimentally (assembled communities), but became increasingly represented in the overall pond metacommunity during the course of the full growing season. In comparison, the composition of experimental mesocosms that lacked any initial zooplankton community (unassembled communities) were found to be driven by dispersal measured at the local pond community but not by dispersal observed across the overall metacommunity. These results suggest that the role of dispersal may shift from a contributor to local, ecological dynamics to that of metacommunity‐wide, colonization–extinction dynamics as communities assemble.