Community assembly is a race between immigration and adaptation: eco‐evolutionary interactions across spatial scales

Both ecological and evolutionary mechanisms have been proposed to describe how natural communities become assembled at both regional and biogeographical scales. Yet, these theories have largely been developed in isolation. Here, we unite these separate views and develop an integrated eco‐evolutionary framework of community assembly. We use a simulation approach to explore the factors determining the interplay between ecological and evolutionary mechanisms systematically across spatial scales. Our results suggest that the same set of ecological and evolutionary processes can determine community assembly at both regional and biogeographical scales. We find that the importance of evolution and community monopolization effects, defined as the eco‐evolutionary dynamics that occur when local adaptation of early established immigrants is fast enough to prevent the later immigration of better pre‐adapted species, are not restricted to adaptive radiations on remote islands. They occur at dispersal rates of up to ten individuals per generation, typical for many species at the scale of regional metacommunities. Dispersal capacity largely determines whether ecological species sorting or evolutionary monopolization structure metacommunity diversity and distribution patterns. However, other factors related to the spatial scale at which community assembly processes are acting, such as metacommunity size and the proportion of empty patches, also affect the relative importance of ecology versus evolution. We show that evolution often determines community assembly, and this conclusion is robust to a wide range of assumptions about spatial scale, mode of reproduction, and environmental structure. Moreover, we found that community monopolization effects occur even though species fully pre‐adapted to each habitat are abundant in the metacommunity, a scenario expected a priori to prevent any meaningful effect of evolution. Our results strongly support the idea that the same eco‐evolutionary processes underlie community assembly at regional and biogeographical scales.     

Metacommunity size influences aquatic community composition in a natural mesocosm landscape

Ecosystems are often arranged in naturally patchy landscapes with habitat patches linked by dispersal of species in a metacommunity. The size of a metacommunity, or number of patches, is predicted to influence community dynamics and therefore the structure and function of local communities. However, such predictions have yet to be experimentally tested using full food webs in natural metacommunities. We used the natural mesocosm system of aquatic macroinvertebrates in bromeliad phytotelmata to test the effect of the number of patches in a metacommunity on species richness, abundance, and community composition. We created metacommunities of varying size using fine mesh cages to enclose a gradient from a single bromeliad up to the full forest. We found that species richness, abundance, and biomass increased from enclosed metacommunities to the full forest size and that diversity and evenness also increased in larger enclosures. Community composition was affected by metacommunity size across the full gradient, with a more even detritivore community in larger metacommunities, and taxonomic groups such as mosquitoes going locally extinct in smaller metacommunities. We were able to divide the effects of metacommunity size into aquatic and terrestrial habitat components and found that the importance of each varied by species; those with simple life cycles were only affected by local aquatic habitat whereas insects with complex life cycles were also affected by the amount of terrestrial matrix. This differential survival of obligate and non‐obligate dispersers allowed us to partition the beta‐diversity between metacommunities among functional groups. Our study is one of the first tests of metacommunity size in a natural metacommunity landscape and shows that both diversity and community composition are significantly affected by metacommunity size. Synthesis Natural food webs are sensitive to meta‐community size, i.e. the number of patches connected through dispersal. We provide an empirical test using the aquatic foodweb associated within bromeliads as a model system. When we reduced the number of bromeliad patches connect through dispersal, we found a clear change of the foodweb in terms of population sizes, beta diversity, community composition and predator‐prey ratios. The response of individual taxa was predictable based on species traits including dispersal modes, life cycle, and adult resource requirements. Our study demonstrates that community structure is strongly influenced by the interplay of species traits and landscape properties.     

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.     

Predicting food‐web structure with metacommunity models

Synthesis Metacommunity theory aims to elucidate the relative influence of local and regional‐scale processes in generating diversity patterns across the landscape. Metacommunity research has focused largely on assemblages of competing organisms within a single trophic level. Here, we test the ability of metacommunity models to predict the network structure of the aquatic food web found in the leaves of the northern pitcher plant Sarracenia purpurea. The species‐sorting and patch‐dynamics models most accurately reproduced nine food web properties, suggesting that local‐scale interactions play an important role in structuring Sarracenia food webs. Our approach can be applied to any well‐resolved food web for which data are available from multiple locations. The metacommunity framework explores the relative influence of local and regional‐scale processes in generating diversity patterns across the landscape. Metacommunity models and empirical studies have focused mostly on assemblages of competing organisms within a single trophic level. Studies of multi‐trophic metacommunities are predominantly restricted to simplified trophic motifs and rarely consider entire food webs. We tested the ability of the patch‐dynamics, species‐sorting, mass‐effects, and neutral metacommunity models, as well as three hybrid models, to reproduce empirical patterns of food web structure and composition in the complex aquatic food web found in the northern pitcher plant Sarracenia purpurea. We used empirical data to determine regional species pools and estimate dispersal probabilities, simulated local food‐web dynamics, dispersed species from regional pools into local food webs at rates based on the assumptions of each metacommunity model, and tested their relative fits to empirical data on food‐web structure. The species‐sorting and patch‐dynamics models most accurately reproduced nine food web properties, suggesting that local‐scale interactions were important in structuring Sarracenia food webs. However, differences in dispersal abilities were also important in models that accurately reproduced empirical food web properties. Although the models were tested using pitcher‐plant food webs, the approach we have developed can be applied to any well‐resolved food web for which data are available from multiple locations.     

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.     

Integrating environmental and spatial processes in ecological community dynamics

The processes controlling the abundances of species across multiple sites form the cornerstone of modern ecology. In these metacommunities, the relative importance of local environmental and regional spatial processes is currently hotly debated, especially in terms of the validity of neutral model. I collected 158 published data sets with information on community structure, environmental and spatial variables. I showed that approximately 50% of the variation in community composition is explained by both environmental and spatial variables. The majority of the data sets were structured by species-sorting dynamics (SS), followed by a combination of SS and mass-effect dynamics. While neutral processes were the only structuring process in 8% of the collected natural communities, disregarding neutral dispersal processes would result in missing important patterns in 37% of the studied communities. Moreover, metacommunity characteristics such as dispersal type, habitat type and spatial scale predicted part of the detected variation in metacommunity structure.     

Disentangling niche and neutral influences on community assembly: assessing the performance of community phylogenetic structure tests

Patterns of phylogenetic relatedness within communities have been widely used to infer the importance of different ecological and evolutionary processes during community assembly, but little is known about the relative ability of community phylogenetics methods and null models to detect the signature of processes such as dispersal, competition and filtering under different models of trait evolution. Using a metacommunity simulation incorporating quantitative models of trait evolution and community assembly, I assessed the performance of different tests that have been used to measure community phylogenetic structure. All tests were sensitive to the relative phylogenetic signal in species metacommunity abundances and traits; methods that were most sensitive to the effects of niche-based processes on community structure were also more likely to find non-random patterns of community phylogenetic structure under dispersal assembly. When used with a null model that maintained species occurrence frequency in random communities, several metrics could detect niche-based assembly when there was strong phylogenetic signal in species traits, when multiple traits were involved in community assembly, and in the presence of environmental heterogeneity. Interpretations of the causes of community phylogenetic structure should be modified to account for the influence of dispersal.     

Metacommunity,mainland‐island system or island communities? Assessing the regional dynamics of plant communities in a fragmented landscape

Understanding the regional dynamics of plant communities is crucial for predicting the response of plant diversity to habitat fragmentation. However, for fragmented landscapes the importance of regional processes, such as seed dispersal among isolated habitat patches, has been controversially debated. Due to the stochasticity and rarity of among‐patch dispersal and colonization events, we still lack a quantitative understanding of the consequences of these processes at the landscape‐scale. In this study, we used extensive field data from a fragmented, semi‐arid landscape in Israel to parameterize a multi‐species incidence‐function model. This model simulates species occupancy pattern based on patch areas and habitat configuration and explicitly considers the locations and the shapes of habitat patches for the derivation of patch connectivity. We implemented an approximate Bayesian computation approach for parameter inference and uncertainty assessment. We tested which of the three types of regional dynamics – the metacommunity, the mainland‐island, or the island communities type – best represents the community dynamics in the study area and applied the simulation model to estimate the extinction debt in the investigated landscape. We found that the regional dynamics in the patch‐matrix study landscape is best represented as a system of highly isolated ‘island’ communities with low rates of propagule exchange among habitat patches and consequently low colonization rates in local communities. Accordingly, the extinction rates in the local communities are the main drivers of community dynamics. Our findings indicate that the landscape carries a significant extinction debt and in model projections 33–60% of all species went extinct within 1000 yr. Our study demonstrates that the combination of dynamic simulation models with field data provides a promising approach for understanding regional community dynamics and for projecting community responses to habitat fragmentation. The approach bears the potential for efficient tests of conservation activities aimed at mitigating future losses of biodiversity.     

Dispersal-mediated trophic interactions can generate apparent patterns of dispersal limitation in aquatic metacommunities

Dispersal is a major organising force in metacommunities, which may facilitate compositional responses of local communities to environmental change and affect ecosystem function. Organism groups differ widely in their dispersal abilities and their communities are therefore expected to have different adaptive abilities. In mesocosms, we studied the simultaneous compositional response of three plankton communities (zoo-, phyto- and bacterioplankton) to a primary productivity gradient and evaluated how this response was mediated by dispersal intensity. Dispersal enhanced responses in all three planktonic groups, which also affected ecosystem functioning. Yet, variation partitioning analyses indicated that responses in phytoplankton and bacterial communities were not only controlled by dispersal directly but also indirectly through complex trophic interactions. Our results indicate that metacommunity patterns emerging from dispersal can cascade through the food web and generate patterns of apparent dispersal limitation in organisms at other trophic levels.     

Evolution in metacommunities: on the relative importance of species sorting and monopolization in structuring communities

Abstract: Adaptive evolution within species and community assembly involving multiple species are both affected by dispersal and spatiotemporal environmental variation and may thus interact with each other. We examined this interaction in a simple three-patch metacommunity and found that these two processes produce very different associations between species composition and local environment. In most conditions, we find a pattern we call "species sorting," wherein local adaptation by resident species cannot prevent invasions by other preadapted species as environmental conditions change (strong association between local environmental conditions and local community composition). When dispersal rates are very low relative to the other two rates, local adaptation by resident species predominates, leading to strong priority effects that prevent successful colonization by other species that would have been well adapted, a pattern we call "local monopolization." When dispersal and evolutionary rates are both very high, we find that an evolving species outcompetes other species in all patches, a pattern we call "global monopolization." When environmental oscillations are very frequent, local monopolization predominates. Our findings indicate that there can be strong modification of community assembly by local adaptive processes and that these depend strongly on the relative rates of evolution, dispersal, and environmental change.     

Gene networks and metacommunities: dispersal differences can override adaptive advantage

Dispersal is an important mechanism contributing to both ecological and evolutionary dynamics. In metapopulation and metacommunity ecology, dispersal enables new patches to be colonized; in evolution, dispersal counter-acts local selection, leading to regional homogenization. Here, I consider a three-patch metacommunity in which two species, each with a limiting quantitative trait underlain by gene networks of 16 to 256 genes, compete with one another and disperse among patches. Incorporating dispersal among heterogeneous patches introduces a tradeoff not observed in single-patch simulations: if the difference between gene network size of the two species is greater than the difference in dispersal ability (e.g., if the ratio of network sizes is larger than the ratio of dispersal abilities), then genetic architecture drives community outcome. However, if the difference in dispersal abilities is greater than gene network differences, then any adaptive advantages afforded by genetic architecture are over-ridden by dispersal. Thus, in addition to the selective pressures imposed by competition that shape the genetic architecture of quantitative traits, dispersal among patches creates an escape that may further alter the effects of different genetic architectures. These results provide a theoretical expectation for what we may observe as the field of ecological genomics develops.     

Consequences of adaptive foraging in diverse communities

1.  Selective pressures acting on foraging activities constrain the strength of interaction, hence the stability and energetic availability in food webs.
2.  Because such selective pressures are usually measured at the individual level and because most experimental and theoretical works focus on simple settings, linking adaptive foraging with community scale patterns is still a far stretch.
3.  Some recent models incorporate foraging adaptation in diverse communities. The models vary in the way they incorporate adaptation, via evolutionary or behavioural changes, and define individual fitness in various ways.
4.  In spite of these differences, some general results linking adaptation to community structure and functioning emerge. In the present article, I introduce these different models and highlight their common results.
5.  Adaptive foraging provides stability to large food web models and predicts successfully interaction patterns within food webs as well as other topological features such as food chain length.
6.  The relationships between adaptive foraging and other structuring factors particularly depend on how well connected the local community is with surrounding communities (metacommunity aspect).     

Determinism of bacterial metacommunity dynamics in the southern East China Sea varies depending on hydrography

Spatial variation of communities composition (metacommunities) results from multiple assembly mechanisms, including environmental filtering and dispersal; however, whether and why the relative importance of the assembly mechanisms in shaping bacterial metacommunity changes through time in marine pelagic systems remains poorly studied. Here, we applied the elements of metacommunity structure framework and the variation partitioning framework to examine whether temporal variation of hydrographic conditions influences bacterioplankton metacommunity dynamics in the southern East China Sea (ECS). The spatiotemporal variation of bacterial communities composition was revealed using 454 pyrosequencing of 16S rDNA. In addition to the whole bacterial community, we analyzed four dominant taxonomic groups (Cyanobacteria, Alphaproteobacteria, Gammaproteobacteria, and Actinobacteria) separately. Our analyses indicate that, considering the whole community level, the determinism of metacommunity structure varied among seasons. When the degree of connectivity was low (December), the metacommunity exhibited random distribution and was explained mainly by the environmental component. However, Clementsian metacommunity was found at intermediate connectivity (May), during which the environmental and spatial predictors were both significant. When connectivity was high (August), a random distribution pattern was found and no significant effect of environmental filtering or dispersal limitation was detected. Nevertheless, when considering different taxonomic groups, the differences in metacommunity dynamics among groups were found. Our results suggest that the driving forces of metacommunity dynamics varied depending on hydrography, as the degrees of environmental heterogeneity and connectivity among habitat patches were determined by circulation pattern. Moreover, mechanisms varied among different taxonomic groups, suggesting that differential dispersal capacity among taxonomic groups should be integrated into community assembly studies.     

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.     

Salinisation reduces biodiversity in neighbouring freshwater patches in experimental metacommunities

1. Aquatic ecosystems are biodiversity hot spots across many landscapes; therefore, the degradation of these habitats can lead to decreases in biodiversity across multiple scales. Salinisation is a global issue that threatens freshwater ecosystems by reducing water quality and local biodiversity. The effects of salinity on local processes have been studied extensively; however, the effects of salinisation or similar environmental stressors within a metacommunity (a dispersal network of several distinct communities) have not been explored.
2. We tested how the spatial heterogeneity and the environmental contrast between freshwater and saline habitat patches influenced cladoceran biodiversity and species composition at local and regional scales in a metacommunity mesocosm experiment. We defined spatial heterogeneity as the proportion of freshwater to saltwater patches within the metacommunity, ranging from a freshwater-dominated metacommunity to a saltwater-dominated metacommunity. Environmental contrast was defined as the environmental distance between habitat patches along the salinity gradient in which low-contrast metacommunities consisted of freshwater and low-salinity patches and high-contrast metacommunities consisted of freshwater and high-salinity patches.
3. We hypothesised that the α-richness of freshwater patches and metacommunity γ-richness would decrease as freshwater patches became less abundant along the spatial heterogeneity gradient in both low- and high-contrast metacommunities, because there would be fewer freshwater patches that could serve as source populations for declining populations. We hypothesised that low-contrast metacommunities would support more species across the spatial heterogeneity gradient than high-contrast metacommunities, because, via dispersal, low-salinity patches can support halotolerant freshwater species that can mitigate population declines in neighbouring freshwater patches, whereas` high-salinity patches will mostly support halophilic species, providing fewer potential colonisers to freshwater patches.
4. We found that α-richness of freshwater mesocosms and metacommunity γ-richness declined in saline-dominated metacommunities regardless of the environmental contrast between the freshwater and saline mesocosms. We found that environmental contrast influenced freshwater and saline community composition in low-contrast metacommunities by increasing the abundances of species that could tolerate low-salinity environments through dispersal, whereas freshwater and high-salinity communities showed limited interactions through dispersal.
5. Freshwater mesocosms had a disproportionate effect on the local and regional biodiversity in these experimental metacommunities, indicating that habitat identity may be more important than habitat diversity for maintaining biodiversity in some metacommunities. This study further emphasises the importance in maintaining multiple species-rich habitat patches across landscapes, particularly those experiencing landscape-wide habitat degradation.

     

Community differentiation on landscapes: drift, migration and speciation

Theories of the differentiation of ecological communities on landscapes have typically not considered evolutionary dynamics. Here we analytically study the expected differentiation among local communities in a large metacommunity, undergoing speciation, ecological drift and intercommunity dispersal, in the context of neutral theory. We demonstrate that heterogeneity in species diversity and abundance arises among communities when local communities are small and intercommunity migration is infrequent. We propose a new measure to describe community differentiation, defined as the average correlation or the average probability (C_st) that two randomly sampled individuals of the same species within local communities are from the same ancestor. The effects of driving forces (migration, mutation, and ecological drift) are incorporated into the two-level hierarchical community structure in a finite island model of neutral communities. Community differentiation can increase the effective metacommunity size or the Hubbell's fundamental species diversity in the metacommunity by a factor (1−C_st)⁻¹. Significant community differentiation arises when C_st≠0. Intercommunity migration promotes species diversity in local communities but reduce species diversity in the metacommunity. In either the finite or infinite island case, one can estimate the number of intercommunity migrants by using multiple local community datasets when the speciation is negligible in the neutral local communities, or by using the metacommunity dataset when the speciation is included in the local neutral communities. These results highlight the significance of the evolutionary mechanisms in generating heterogeneous communities in the absence of complicated ecological processes on large landscapes.     

Network isolation and local diversity in neutral metacommunities

Biologists seek an understanding of the biological and environmental factors determining local community diversity. Recent advances in metacommunity ecology, and neutral theory in particular, highlight the importance of dispersal processes interacting with the spatial structure of a landscape for generating spatial patterns and maintaining biodiversity. The relative spatial isolation of a community is traditionally thought to have a large influence on local diversity. However, isolation remains an elusive concept to quantify, particularly in metacommunities with complex spatial structure. We represent the metacommunity as a network of local communities, and use network centrality measures to quantify the isolation of a local community. Using spatially explicit neutral theory, we examine how node position predicts variation in alpha diversity across a metacommunity. We find that diversity increases with node centrality in the network, but only when centrality is measured on a given scale in the network that widens with increasing dispersal rates and narrows with increasing evolutionary rates. More generally, complex biodiversity patterns form only when the underlying geography has structure on this critical scale. This provides a framework for understanding the influence of spatial geographic structure on global biodiversity patterns.     

The importance of landscape and habitat properties in explaining instantaneous and long‐term distributions of large branchiopods in subtropical temporary pans

1. The notion that the spatial configuration of habitat patches has to be taken into account to understand the structure and dynamics of ecological communities is the starting point of metacommunity ecology. One way to assess metacommunity structure is to investigate the relative importance of environmental heterogeneity and spatial structure in explaining community patterns over different spatial and temporal scales. 2. We studied metacommunity structure of large branchiopod assemblages characteristic of subtropical temporary pans in SE Zimbabwe using two community data sets: a community snapshot and a long‐term data set covering 4 years. We assessed the relative importance of environmental heterogeneity and dispersal (inferred from patch occupancy patterns) as drivers of community structure. Furthermore, we contrasted metacommunity patterns in pans that occasionally connect to the river (floodplain pans) and pans that lack such connections altogether (endorheic pans) using redundancy models. 3. Echoes of species sorting and dispersal limitation emerge from our data set, suggesting that both local and regional processes contribute to explaining branchiopod assemblages in this system. Relative importance of local and regional factors depended on the type of data set considered. Overall, habitat characteristics that vary in time, such as conductivity, hydroperiod and vegetation cover, best explained the instantaneous species composition observed during a snapshot sampling while long‐term species composition appeared to be linked to more constant intrinsic habitat properties such as river connectivity and spatial location.     

Any way the wind blows - frequent wind dispersal drives species sorting in ephemeral aquatic communities

Despite an upsurge of interest in spatial interactions between communities and in the impact of dispersal on ecological and evolutionary processes, dispersal patterns and dynamics in natural metacommunities remain poorly understood. Although passive aerial dispersal of freshwater invertebrates is generally accepted, the frequency and relative importance of wind as a vector is still subject of considerable debate. We assessed the importance of wind dispersal in an invertebrate metacommunity in a cluster of 36 temporary rock pools on an isolated mountaintop in South Africa. Wind dispersal was quantified every four days using nine windsocks (about 1.5 m above rock base), placed in the field during one month. Distance to the nearest pool varied from 2 up to 16 m. Wind direction and speed were monitored for the entire period. About 850 propagules (mostly resting eggs) of 17 taxa were captured. The presence of water in the pools (level of exposure of the dormant propagule bank) and the dominant wind direction were the key factors affecting the yield. Wind speed was much less important.
Our results suggest that wind dispersal of propagules from temporary aquatic systems is more frequent than previously thought. This may stabilise the metacommunity by mediating gene flow among populations and facilitating rapid (re)colonisation of patches. On the other hand, wind erosion of the dormant propagule bank may lead to egg bank depletion and local extinction.
The measured frequent wind dispersal most likely fuels strong species sorting processes ultimately shaping the structure of the local communities as observed in an earlier study. To elucidate the link between local dispersal rates and their contribution to long range dispersal is a major challenge for future research on aerial dispersal of aquatic invertebrates.     

Positive interactions and the emergence of community structure in metacommunities

The significant role of space in maintaining species coexistence and determining community structure and function is well established. However, community ecology studies have mainly focused on simple competition and predation systems, and the relative impact of positive interspecific interactions in shaping communities in a spatial context is not well understood. Here we employ a spatially explicit metacommunity model to investigate the effect of local dispersal on the structure and function of communities in which species are linked through an interaction web comprising mutualism, competition and exploitation. Our results show that function, diversity and interspecific interactions of locally linked communities undergo a phase transition with changes in the rate of species dispersal. We find that low spatial interconnectedness favors the spontaneous emergence of strongly mutualistic communities which are more stable but less productive and diverse. On the other hand, high spatial interconnectedness promotes local biodiversity at the expense of local stability and supports communities with a wide range of interspecific interactions. We argue that investigations of the relationship between spatial processes and the self-organization of complex interaction webs are critical to understanding the geographic structure of interactions in real landscapes.