The metacommunity concept: a framework for multi-scale community ecology

The metacommunity concept is an important way to think about linkages between different spatial scales in ecology. Here we review current understanding about this concept. We first investigate issues related to its definition as a set of local communities that are linked by dispersal of multiple potentially interacting species. We then identify four paradigms for metacommunities: the patch‐dynamic view, the species‐sorting view, the mass effects view and the neutral view, that each emphasizes different processes of potential importance in metacommunities. These have somewhat distinct intellectual histories and we discuss elements related to their potential future synthesis. We then use this framework to discuss why the concept is useful in modifying existing ecological thinking and illustrate this with a number of both theoretical and empirical examples. As ecologists strive to understand increasingly complex mechanisms and strive to work across multiple scales of spatio‐temporal organization, concepts like the metacommunity can provide important insights that frequently contrast with those that would be obtained with more conventional approaches based on local communities alone.     

Metacommunity organisation,spatial extent and dispersal in aquatic systems: patterns,processes and prospects

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Linking community and ecosystem dynamics through spatial ecology

Classical approaches to food webs focus on patterns and processes occurring at the community level rather than at the broader ecosystem scale, and often ignore spatial aspects of the dynamics. However, recent research suggests that spatial processes influence both food web and ecosystem dynamics, and has led to the idea of 'metaecosystems'. However, these processes have been tackled separately by 'food web metacommunity' ecology, which focuses on the movement of traits, and 'landscape ecosystem' ecology, which focuses on the movement of materials among ecosystems. Here, we argue that this conceptual gap must be bridged to fully understand ecosystem dynamics because many natural cases demonstrate the existence of interactions between the movements of traits and materials. This unification of concepts can be achieved under the metaecosystem framework, and we present two models that highlight how this framework yields novel insights. We then discuss patches, limiting factors and spatial explicitness as key issues to advance metaecosystem theory. We point out future avenues for research on metaecosystem theory and their potential for application to biological conservation.     

Experimental evidence that soil heterogeneity enhances plant diversity during community assembly

Aims Environmental heterogeneity is a primary mechanism explaining species coexistence and extant patterns of diversity. Despite strong theoretical support and ample observational evidence, few experimental studies in plant communities have been able to demonstrate a causal link between environmental heterogeneity and plant diversity. This lack of experimental evidence suggests that either fine-scale heterogeneity has weak effects on plant diversity or previous experiments have been unable to effectively manipulate heterogeneity. Here, we utilize a unique soil manipulation to test whether fine-scale soil heterogeneity will increase plant richness through species sorting among experimental patch types.Methods This experiment was conducted in the tallgrass prairie region of south-central Kansas, USA. We utilized the inherent variation found in the vertical soil profile, which varied in both biotic and abiotic characteristics, and redistributed these strata into either homogeneous or heterogeneous spatial arrangements in 2.4×2.4 m plots. After the soil manipulation, 34 native prairie species were sown into all plots. We conducted annual censuses at peak biomass to quantify species composition and plant density by species within the experimental communities.Important findings After 2 years, species richness was significantly higher in heterogeneous relative to homogeneous plots and this pattern was independent of total plant density. In the heterogeneous plots, 13 species had higher establishment in a specific patch type representing one of the three soil strata. Conversely, no species had greater establishment in the mixed stratum, which comprised the homogeneous plots, relative to the heterogeneous strata. These species sorting patterns suggest that fine-scale heterogeneity creates opportunities for plant establishment due to niche differences, which translates into increased plant diversity at the plot scale. Species richness was more strongly related to plant density among patches comprising homogenous plots—where fine-scale heterogeneity was minimized, but weak in heterogeneous plots. This pattern is consistent with the idea that richness–density relationships dominate when neutral processes are important but are weak when niche processes operate. Unlike many previous attempts, our results provide clear, experimental evidence that fine-scale soil heterogeneity increases species richness through species sorting during community assembly.     

Control of stream insect assemblages: roles of spatial configuration and local environmental factors

Abstract 1. Current views in ecology emphasise that community structure is the sum of multiple processes, with imprints of both regional and local drivers. However, the degree to which stream insect assemblages are structured by spatial configuration (complying with the dispersal‐based neutral hypothesis) and local environmental features (complying with the niche‐based species sorting hypothesis) has not been rigorously examined based on surveys in multiple years. 2. Stream sites in a boreal drainage system were surveyed during three consecutive years and the relative contribution of spatial configuration and local environmental variables to aquatic insect assemblage structure (characterised by both abundance and presence–absence data) was assessed. Separate analyses were conducted for mayflies (Ephemeroptera), stoneflies (Plecoptera), caddisflies (Trichoptera), and non‐biting midges (Diptera: Chironomidae) in each year. 3. There were no relationships between the spatial location and local environmental features of streams in Mantel tests, facilitating exploration of their independent effects on assemblage structure. The study found virtually no effects of spatial location on stream insect assemblages across the study drainage system, as evidenced by Mantel tests and canonical correspondence analyses (CCA). The environmental variables were also rather weakly associated with assemblage structure, with the total amount of explained variation ranging from 9.8% to 31.7% in the CCAs. There were no appreciable differences in the amount of environment‐related explained variation in assemblage structure between mayflies, stoneflies, caddisflies, and midges, but some between‐year differences were noticeable in most insect groups. The environmental variables that were significantly related to assemblage structure exhibited some between‐group and between‐year variability. In general, patterns shown by abundance and presence–absence data were highly similar. 4. It appears that stream insect assemblages comply with the niche‐based species sorting hypothesis in the context of metacommunity ecology. In contrast, the absence of spatial structuring suggests that stream insect assemblages do not comply with the neutral hypothesis, being not strongly dispersal limited at the within‐drainage basin scale.     

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.     

The relative importance of dispersal and local processes in structuring phytoplankton communities in a set of highly interconnected ponds

1. The recognition that both local and regional processes act together in shaping local communities makes determining their relative roles in natural communities central to understanding patterns in community structure. 2. We investigated the relative influence of these processes on the phytoplankton communities of a highly interconnected pond system. We sampled the phytoplankton communities of 28 ponds concurrently with 20 local environmental variables. 3. We found that phytoplankton community variation, in terms of both phytoplankton community composition (PCC) and diversity, was only significantly explained by local environmental variables. These were mainly associated with the contrasting clear‐water and turbid ecological states of the shallow ponds studied. Clear‐water conditions favoured only a few taxa, resulting in a significantly lower taxon diversity and richness under these conditions. 4. The failure to explain variation in PCC by a dispersal model based on the water flow between ponds points at very effective species sorting. This is attributed to the high population turn‐over rates and sensitivity to environmental conditions of phytoplankton communities. Some evidence was found, however, that dispersal influences local communities through mass effects between neighbouring ponds. 5. Overall, our results emphasize both the strong selection pressure that components of the food web exert on phytoplankton communities and the high potential of these communities to respond to such environmental change, thereby effectively opposing the homogenizing effects of continuous dispersal.     

Local community size mediates ecological drift and competition in metacommunities

The outcome of competitive interactions is likely to be influenced by both competitive dominance (i.e. niche-based dynamics) and ecological drift (i.e. neutral dynamics governed by demographic stochasticity). However, spatial models of competition rarely consider the joint operation of these two processes. We develop a model based on the original competition-colonization trade-off model that incorporates niche and neutral processes and several realistic facets of ecological dynamics: it allows local competition (i.e. competition within a patch) to occur within communities of a finite size, it allows competitors to vary in the degree of competitive asymmetry, and it includes the role of local migration (i.e. propagule pressure). The model highlights the role of community size, i.e. the number of competitors in the local community, in mediating the relative importance of stochastic and deterministic forces. In metacommunities where local communities are small, ecological drift is substantial enough that strong competitors become effectively neutral, creating abrupt changes in the outcome of competition not predicted by the standard competition-colonization trade-off. Importantly, the model illustrates that, even when other aspects of species interactions (e.g. migration ability, competitive ability) are unchanged, local community size can alter the dynamics of metacommunity persistence. Our work demonstrates that activities which reduce the size of local communities, such as habitat destruction and degradation, effectively compound the extinction debt.     

The importance of metacommunity ecology for environmental assessment research in the freshwater realm

Most bioassessment programs rest on the assumption that species have different niches, and that abiotic environmental conditions and changes therein determine community structure. This assumption is thus equivalent to the species sorting perspective (i.e. that species differ in their responses to environmental variation) in metacommunity ecology. The degree to which basing bioassessment on the species sorting perspective is reasonable is likely to be related to the spatial extent of a study and the characteristics of the organism groups (e.g. dispersal ability) with which the effects of anthropogenic changes are assessed. Recent findings in metacommunity research have stressed that community structure is determined not only by local abiotic environmental conditions but also by biotic interactions and dispersal‐related effects. For example, dispersal limitation may prevent community structure recovery from the effects of a putative stressor, as organisms may not be able to disperse to all sites in a region. Mass effects (i.e. the presence of species in environmentally suboptimal sites due to high dispersal rates from environmentally suitable sites) may, in turn, obscure the effects of a stressor, as dispersal from source sites (e.g. an unaltered site) allows persistence at sink sites (e.g. an anthropogenically altered site). Better bioassessment should thus take both niche‐ and dispersal‐related processes simultaneously into consideration, which can be accomplished by explicitly modelling spatial location as a proxy for dispersal effects. Such an integrated approach should be included in bioassessment programs using general multivariate approaches, predictive modelling, and multimetric indices.     

Environmental and spatial processes influencing phytoplankton biomass along a reservoirs‐river‐floodplain lakes gradient: A metacommunity approach

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Maternal effects provide phenotypic adaptation to local environmental conditions

In outcrossing plants, seed dispersal distance is often less than pollen movement. If the scale of environmental heterogeneity within a population is greater than typical seed dispersal distances but less than pollen movement, an individual's environment will be similar to that of its mother but not necessarily its father. Under these conditions, environmental maternal effects may evolve as a source of adaptive plasticity between generations, enhancing offspring fitness in the environment that they are likely to experience. This idea is illustrated using Campanula americana, an herb that grows in understory and light-gap habitats. Estimates of seed dispersal suggest that offspring typically experience the same light environment as their mother. In a field experiment testing the effect of open vs understory maternal light environments, maternal light directly influenced offspring germination rate and season, and indirectly affected germination season by altering maternal flowering time. Results to date indicate that these maternal effects are adaptive; further experimental tests are ongoing. Evaluating maternal environmental effects in an ecological context demonstrates that they may provide phenotypic adaptation to local environmental conditions.     

Spatial coherence and the persistence of high diversity in spatially heterogeneous landscapes

Our planet hosts a variety of highly diverse ecosystems. The persistence of high diversity is generally attributed to factors such as the structure of interactions among species and the dispersal of species in metacommunities. Here, we show that large contiguous landscapes—that are characterized by high dispersal—facilitate high species richness due to the spatial heterogeneity in interspecies interactions. We base our analysis on metacommunities under high dispersal where species densities become equal across habitats (spatially coherent). We find that the spatially coherent metacommunity can be represented by an effective species interaction‐web that has a significantly lower complexity than the constituent habitats. Our framework also explains how spatial heterogeneity eliminates differences in the effective interaction‐web, providing a basis for deviations from the area‐heterogeneity tradeoff. These results highlight the often‐overlooked case of high dispersal where spatial coherence provides a novel mechanism for supporting high diversity in large heterogeneous landscapes.     

Neutral community dynamics, the mid-domain effect and spatial patterns in species richness

The mid‐domain effect (MDE) aims to explain spatial patterns in species richness invoking only stochasticity and geometrical constraints. In this paper, we used simulations to show that its main qualitative prediction, a hump‐shaped pattern in species richness, converges to the expectation of a spatially bounded neutral model when communities are linked by short‐distance migration. As these two models can be linked under specific situations, neutral theory may provide a mechanistic population level basis for MDE. This link also allows establishing in which situations MDE patterns are more likely to be found. Also, in this situation, MDE models could be used as a first approximation to understand the role of both stochastic (ecological drift and migration) and deterministic (adaptation to environmental conditions) processes driving the spatial structure of species richness.     

Beta‐diversity in temperate and tropical forests reflects dissimilar mechanisms of community assembly

Site‐to‐site variation in species composition (β‐diversity) generally increases from low‐ to high‐diversity regions. Although biogeographical differences in community assembly mechanisms may explain this pattern, random sampling effects can create this pattern through differences in regional species pools. Here, we compared assembly mechanisms between spatially extensive networks of temperate and tropical forest plots with highly divergent species pools (46 vs. 607 species). After controlling for sampling effects, β‐diversity of woody plants was similar and higher than expected by chance in both forests, reflecting strong intraspecific aggregation. However, different mechanisms appeared to explain aggregation in the two forests. In the temperate forest, aggregation reflected stronger environmental correlations, suggesting an important role for species‐sorting (e.g. environmental filtering) processes, whereas in the tropics, aggregation reflected stronger spatial correlations, more likely reflecting dispersal limitation. We suggest that biogeographical differences in the relative importance of different community assembly mechanisms contribute to these striking gradients in global biodiversity.     

Environmental and spatial controls of biotic assemblages in a discrete semi‐terrestrial habitat: comparison of organisms with different dispersal abilities sampled in the same plots

Aim The development of metacommunity theory inspired a series of studies exploring the importance of environmental and spatial effects on the composition of biotic assemblages. However, the comparison of different groups of organisms has been hampered by differences in sampling design, spatial scales or the environmental variables involved. Our aim was to test how dispersal ability affects metacommunity structure and associated species distributions by sampling different species groups in the same plots to avoid these problems. Location Western Carpathian Mountains (Europe). Methods In 191 fens we sampled the composition of diatom, bryophyte, vascular plant and mollusc assemblages, water chemistry, and macroclimatic data. We then generated spatial variables covering all relevant spatial scales using analysis of principal coordinates of neighbour matrices (PCNM). We applied the adjusted variation partitioning algorithm to quantify the effects of environment and space. Results Pure effects of water chemistry and space were highly significant for all groups of organisms. Spatial effects were stronger for groups with larger propagules (vascular plants, molluscs) than for those with smaller propagules (diatoms, bryophytes). Assemblages of macroscopic bryophytes were structured slightly less by geography and much more by environment than were those of microscopic diatoms. Vascular plant and mollusc assemblages turned out to be more spatially structured (as compared to diatom and bryophyte assemblages), with small differences between the two groups. Coarse‐scale spatial effects dominated in the bryophyte metacommunity, while in the other groups, including diatoms, finer‐scale effects were also important. Main conclusions Given that our analyses are based on a standardized sampling and analytical framework, our findings provide strong support for the hypothesis that both environmental and spatial variables structure metacommunities of organisms with very different dispersal abilities, including microscopic diatoms. In addition, we show for the first time that the strengths of these effects and their scale dependence may be predicted using important trait differences between organisms, for example differences in propagule size.