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1.
1. Ecologists have recognised several factors that may explain the distribution of species in a metacommunity. These factors may be related to the dispersal of individuals among the patches and environmental conditions. 2. Here, we attempted to determine which of the four different metacommunity frameworks (patch dynamics, mass effect, neutral processes, and species sorting) explain the distribution of Arctiinae moths in Brazilian savanna areas with different tree species richness. 3. The Arctiinae moths were categorised as habitat specialists or generalists, common or rare, and belonging to the tribes Arctiini and Lithosiini. We hypothesized that environmental variables best explain the abundance and occurrence of habitat specialist species, common species, and members of Lithosiini; whereas spatial processes are more closely related to habitat generalists, rare species, and members of Arctiini. 4. Contrary to our expectations, we found that the species sorting (mainly dictated by the species richness of trees) best explained the variation in abundance and occurrence of the majority of species groups. Spatial processes (more related to patch dynamics, mass effect, and neutral), although they were significantly related to some species groups, were not strong enough to explain the distribution of these species in the study area. 5. The plant species richness was the most important environmental condition, related to moth species niches. Therefore, species sorting best explained the distribution of the species of Arctiinae in the Brazilian savanna.  相似文献   

2.
Niche and neutral processes drive community assembly and metacommunity dynamics, but their relative importance might vary with the spatial scale. The contribution of niche processes is generally expected to increase with increasing spatial extent at a higher rate than that of neutral processes. However, the extent to what community composition is limited by dispersal (usually considered a neutral process) over increasing spatial scales might depend on the dispersal capacity of composing species. To investigate the mechanisms underlying the distribution and diversity of species known to have great powers of dispersal (hundreds of kilometres), we analysed the relative importance of niche processes and dispersal limitation in determining beta‐diversity patterns of aquatic plants and cladocerans over regional (up to 300 km) and continental (up to 3300 km) scales. Both taxonomic groups were surveyed in five different European regions and presented extremely high levels of beta‐diversity, both within and among regions. High beta‐diversity was primarily explained by species replacement (turnover) rather than differences in species richness (i.e. nestedness). Abiotic and biotic variables were the main drivers of community composition. Within some regions, small‐scale connectivity and the spatial configuration of sampled communities explained a significant, though smaller, fraction of compositional variation, particularly for aquatic plants. At continental scale (among regions), a significant fraction of compositional variation was explained by a combination of spatial effects (exclusive contribution of regions) and regionally‐structured environmental variables. Our results suggest that, although dispersal limitation might affect species composition in some regions, aquatic plant and cladoceran communities are not generally limited by dispersal at the regional scale (up to 300 km). Species sorting mediated by environmental variation might explain the high species turnover of aquatic plants and cladocerans at regional scale, while biogeographic processes enhanced by dispersal limitation among regions might determine the composition of regional biotas.  相似文献   

3.
Two‐patch compartment models have been explored to understand the spatial processes that promote species coexistence. However, a phenomenological definition of the inter‐patch ‘dispersal rate’ has limited the quantitative predictability of these models to community dynamics in spatially continuous habitats. Here, we mechanistically rederived a two‐patch Lotka–Volterra competition model for a spatially continuous reaction‐diffusion system where a narrow corridor connects two large habitats. We provide a mathematical formula of the dispersal rate appearing in the two‐patch compartment model as a function of habitat size, corridor shape (ratio of its width to its length), and organism diffusion coefficients. For most reasonable settings, the two‐patch compartment model successfully approximated not only the steady states, but also the transient dynamics of the reaction–diffusion model. Further numerical simulations indicated the general applicability of our formula to other types of community dynamics, e.g. driven by resource‐competition, in spatially homogeneous and heterogeneous environments. Our results suggest that the spatial configuration of habitats plays a central role in community dynamics in space. Furthermore, our new framework will help to improve experimental designs for quantitative test of metacommunity theories and reduce the gaps among modeling, empirical studies, and their application to landscape management.  相似文献   

4.
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.  相似文献   

5.
6.
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.  相似文献   

7.
The correspondence between species distribution and the environment depends on species’ ability to track favorable environmental conditions (via dispersal) and to maintain competitive hierarchy against the constant influx of migrants (mass effect) and demographic stochasticity (ecological drift). Here we report a simulation study of the influence of landscape structure on species distribution. We consider lottery competition for space in a spatially heterogeneous environment, where the landscape is represented as a network of localities connected by dispersal. We quantified the contribution of neutrality and species sorting to their spatial distribution. We found that neutrality increases and the strength of species-sorting decreases with the centrality of a community in the landscape when the average dispersal among communities is low, whereas the opposite was found at elevated dispersal. We also found that the strength of species-sorting increases with environmental heterogeneity. Our results illustrate that spatial structure of the environment and of dispersal must be taken into account for understanding species distribution. We stress the importance of spatial geographic structure on the relative importance of niche vs. neutral processes in controlling community dynamics.  相似文献   

8.
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.  相似文献   

9.
Patch occupancy theory predicts that a trade-off between competition and dispersal should lead to regional coexistence of competing species. Empirical investigations, however, find local coexistence of superior and inferior competitors, an outcome that cannot be explained within the patch occupancy framework because of the decoupling of local and spatial dynamics. We develop two-patch metapopulation models that explicitly consider the interaction between competition and dispersal. We show that a dispersal-competition trade-off can lead to local coexistence provided the inferior competitor is superior at colonizing empty patches as well as immigrating among occupied patches. Immigration from patches that the superior competitor cannot colonize rescues the inferior competitor from extinction in patches that both species colonize. Too much immigration, however, can be detrimental to coexistence. When competitive asymmetry between species is high, local coexistence is possible only if the dispersal rate of the inferior competitor occurs below a critical threshold. If competing species have comparable colonization abilities and the environment is otherwise spatially homogeneous, a superior ability to immigrate among occupied patches cannot prevent exclusion of the inferior competitor. If, however, biotic or abiotic factors create spatial heterogeneity in competitive rankings across the landscape, local coexistence can occur even in the absence of a dispersal-competition trade-off. In fact, coexistence requires that the dispersal rate of the overall inferior competitor not exceed a critical threshold. Explicit consideration of how dispersal modifies local competitive interactions shifts the focus from the patch occupancy approach with its emphasis on extinction-colonization dynamics to the realm of source-sink dynamics. The key to coexistence in this framework is spatial variance in fitness. Unlike in the patch occupancy framework, high rates of dispersal can undermine coexistence, and hence diversity, by reducing spatial variance in fitness.  相似文献   

10.
11.
A key challenge for community ecology is to understand to what extent observational data can be used to infer the underlying community assembly processes. As different processes can lead to similar or even identical patterns, statistical analyses of non‐manipulative observational data never yield undisputable causal inference on the underlying processes. Still, most empirical studies in community ecology are based on observational data, and hence understanding under which circumstances such data can shed light on assembly processes is a central concern for community ecologists. We simulated a spatial agent‐based model that generates variation in metacommunity dynamics across multiple axes, including the four classic metacommunity paradigms as special cases. We further simulated a virtual ecologist who analysed snapshot data sampled from the simulations using eighteen output metrics derived from beta‐diversity and habitat variation indices, variation partitioning and joint species distribution modelling. Our results indicated two main axes of variation in the output metrics. The first axis of variation described whether the landscape has patchy or continuous variation, and thus was essentially independent of the properties of the species community. The second axis of variation related to the level of predictability of the metacommunity. The most predictable communities were niche‐based metacommunities inhabiting static landscapes with marked environmental heterogeneity, such as metacommunities following the species sorting paradigm or the mass effects paradigm. The most unpredictable communities were neutral‐based metacommunities inhabiting dynamics landscapes with little spatial heterogeneity, such as metacommunities following the neutral or patch sorting paradigms. The output metrics from joint species distribution modelling yielded generally the highest resolution to disentangle among the simulated scenarios. Yet, the different types of statistical approaches utilized in this study carried complementary information, and thus our results suggest that the most comprehensive evaluation of metacommunity structure can be obtained by combining them.  相似文献   

12.
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.  相似文献   

13.
1. Quantifying the relative importance of environmental filtering versus regional spatial structuring has become an intensively studied area in the context of metacommunity ecology. However, most studies have evaluated the role of environmental and spatial processes using taxonomic data sets of single snapshot surveys. 2. Here, we examined temporal changes in patterns and possible processes behind the functional metacommunity organization of stream fishes in a human‐modified landscape. Specifically, we (i) studied general changes in the functional composition of fish assemblages among 40 wadeable stream sites during a 3‐year study period in the catchment area of Lake Balaton, Hungary, (ii) quantified the relative importance of spatial and environmental factors as determinants of metacommunity structure and (iii) examined temporal variability in the relative role of spatial and environmental processes for this metacommunity. 3. Partial triadic analysis showed that assemblages could be effectively ordered along a functional gradient from invertebrate consuming species dominated by the opportunistic life‐history strategy, to assemblages with a diverse array of functional attributes. The analysis also revealed that functional fish assemblage structure was moderately stable among the sites between the sampling periods. 4. Despite moderate stability, variance partitioning using redundancy analyses (RDA) showed considerable temporal variability in the contribution of environmental and spatial factors to this pattern. The analyses also showed that environmental variables were, in general, more important than spatial ones in determining metacommunity structure. Of these, natural environmental variables (e.g. altitude, velocity) proved to be more influential than human‐related effects (e.g. pond area, % inhabited area above the site, nutrient enrichment), even in this landscape with relatively low variation in altitude and stream size. 5. Pond area was, however, the most important human stressor variable that was positively associated with the abundance of non‐native species with diverse functional attributes. The temporal variability in the relative importance of environmental and spatial factors was probably shaped by the release of non‐native fish from fish ponds to the stream system during flood events. 6. To conclude, both spatial processes and environmental control shape the functional metacommunity organization of stream fish assemblages in human‐modified landscapes, but their importance can vary in time. We argue, therefore, that metacommunity studies should better consider temporal variability in the ecological mechanisms (e.g. dispersal limitation, species sorting) that determine the dynamics of landscape‐level community organization.  相似文献   

14.
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.  相似文献   

15.
The relative importance of niche requirements and dispersal limitation in controlling the landscape‐scale distribution of plants is still contentious. Local occurrence and abundance of alpine plants are commonly thought to be driven by abiotic site conditions due to pronounced environmental gradients over short distances. However, explicit tests of the additional role of dispersal‐related processes for alpine plant distribution patterns are lacking. Here, we combine niche‐based species distribution models with variables describing patch size and connectivity to evaluate if, besides abiotic limitations, spatial habitat configuration affects the occurrence and abundance of six plant species inhabiting patchy snowbed mosaics of the northeastern Calcareous Alps in Austria. Moreover, we assess if eventual effects of spatial patch configuration are more clearly detectable when calculating connectivity based on parameterized mechanistic dispersal kernels for both wind and animal vectors instead of using nearest neighbour metrics. We show that patch size and connectivity are significantly correlated to the occurrence of all and to the abundance of four out of six study species, although the relative importance of these variables, as compared to niche constraints, varies among species. In addition, connectivity measures derived from parameterized dispersal kernels were more closely related to occupancy, and in particular to abundance patterns than a simple nearest neighbour metric. The fitted kernels also suggest that dispersal by alpine chamois plays an important role for inter‐patch seed exchange. We conclude that, despite evident abiotic limitations, recurrent local extinctions and delayed re‐colonizations indeed play a role for the distribution of our study species, and that alpine plants may hence be less in equilibrium with their abiotic environment than commonly thought. Moreover, the relatively high long‐distance dispersal probabilities of animal kernels indicate that the ability of alpine plants to adapt their ranges to a rapidly warming climate may, among other factors, depend on the availability of dispersal services by large mammals.  相似文献   

16.
The metacommunity concept: a framework for multi-scale community ecology   总被引:13,自引:3,他引:13  
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.  相似文献   

17.
阿拉善荒漠啮齿动物集合群落实证研究   总被引:3,自引:2,他引:1  
当生态学家探求在破碎化的栖息地中,群落物种的共存机制、多样性、局域尺度的性质和过程被放到更广阔的时空框架内时,就出现了"集合群落"这一概念。Leibold提出了集合群落概念,他们将一个集合群落定义为局域群落集,这些群落由各个潜在的相互作用的物种的扩散连接在一起。集合群落理论描述了那些发生在集合群落尺度上的过程,并且提出思考关于物种相互作用的新方法。集合群落概念为群落生态学提供了一个新的革命性的范式,集合群落研究的最基本问题是同一系统中多物种共存的机理、多样性的形成原因与维持机制。该范式强调区域范围内群落中的综合变异,强调环境特证和栖息地之间通过扩散调节的生物相互作用和空间变化。Leibold等提出了解释集合群落结果理论上的4个生态范式,即(1)中性理论;(2)斑块动态理论;(3)物种分配理论;(4)集团效应理论。之后有大量有关检验这4种生态理论的研究,但是有关陆地脊椎动物系统的集合群落的研究较少。2010—2012年,通过在内蒙古阿拉善荒漠景观中的8个固定样地中,对啮齿动物、栖息地环境因子进行调查。利用冗余分析和偏冗余分析,评估环境特征和空间特征对物种组成的影响。结果表明,环境特征独自解释72.8%的啮齿动物物种组成变化,空间特征独自解释33.8%的物种组成变化,环境特征和空间特征共同解释86.5%的啮齿动物物种组成变化,结果显著(P=0.032);去除环境特征之后,空间特征解释13.7%的变化(P=0.246),结果不显著;去除空间特征之后,栖息地变化解释52.7%的变化(P=0.016);环境特征和空间特征的交互作用解释20.1%的物种组成的变化,该区域啮齿动物群落构成集合群落,物种共存中环境特征起着主导作用,由物种分配理论解释该集合群落结构。  相似文献   

18.
Dispersal within metacommunities can play a major role in species persistence by promoting asynchrony between communities. Understanding this role is crucial both for explaining species coexistence and managing landscapes that are increasingly fragmented by human activities. Here, we demonstrate that spatial patterning of dispersal connections can drastically alter both the tendency toward asynchrony and the effect of asynchrony on metacommunity dynamics commonly used to infer the potential for persistence. We also demonstrate that changes in dispersal connections in strictly homogeneous predator-prey metacommunities can generate an extremely rich variety of dynamics even when previously investigated properties of connectivity such as the magnitude and distribution of dispersal among patches are held constant. Furthermore, the dynamics we observe depend strongly on initial conditions. Our results illustrate the effectiveness of measures of spatial structure for predicting asynchrony and its effects on community dynamics, providing a deeper understanding of the relationship between spatial structure and species persistence in metacommunities.  相似文献   

19.
Lakes and their topological distribution across Earth's surface impose ecological and evolutionary constraints on aquatic metacommunities. In this study, we group similar lake ecosystems as metacommunity units influencing diatom community structure. We assembled a database of 195 lakes from the tropical Andes and adjacent lowlands (8°N–30°S and 58–79°W) with associated environmental predictors to examine diatom metacommunity patterns at two different levels: taxon and functional (deconstructed species matrix by ecological guilds). We also derived spatial variables that inherently assessed the relative role of dispersal. Using complementary multivariate statistical techniques (principal component analysis, cluster analysis, nonmetric multidimensional scaling, Procrustes, variance partitioning), we examined diatom–environment relationships among different lake habitats (sediment surface, periphyton, and plankton) and partitioned community variation to evaluate the influence of niche‐ and dispersal‐based assembly processes in diatom metacommunity structure across lake clusters. The results showed a significant association between geographic clusters of lakes based on gradients of climate and landscape configuration and diatom assemblages. Six lake clusters distributed along a latitudinal gradient were identified as functional metacommunity units for diatom communities. Variance partitioning revealed that dispersal mechanisms were a major contributor to diatom metacommunity structure, but in a highly context‐dependent fashion across lake clusters. In the Andean Altiplano and adjacent lowlands of Bolivia, diatom metacommunities are niche assembled but constrained by either dispersal limitation or mass effects, resulting from area, environmental heterogeneity, and ecological guild relationships. Topographic heterogeneity played an important role in structuring planktic diatom metacommunities. We emphasize the value of a guild‐based metacommunity model linked to dispersal for elucidating mechanisms underlying latitudinal gradients in distribution. Our findings reveal the importance of shifts in ecological drivers across climatic and physiographically distinct lake clusters, providing a basis for comparison of broad‐scale community gradients in lake‐rich regions elsewhere. This may help guide future research to explore evolutionary constraints on the rich Neotropical benthic diatom species pool.  相似文献   

20.
Models of metapopulations have often ignored local community dynamics and spatial heterogeneity among patches. However, persistence of a community as a whole depends both on the local interactions and the rates of dispersal between patches. We study a mathematical model of a metacommunity with two consumers exploiting a resource in a habitat of two different patches. They are the exploitative competitors or the competing predators indirectly competing through depletion of the shared resource. We show that they can potentially coexist, even if one species is sufficiently inferior to be driven extinct in both patches in isolation, when these patches are connected through diffusive dispersal. Thus, dispersal can mediate coexistence of competitors, even if both patches are local sinks for one species because of the interactions with the other species. The spatial asynchrony and the competition-colonization trade-off are usual mechanisms to facilitate regional coexistence. However, in our case, two consumers can coexist either in synchronous oscillation between patches or in equilibrium. The higher dispersal rate of the superior prompts rather than suppresses the inferior. Since differences in the carrying capacity between two patches generate flows from the more productive patch to the less productive, loss of the superior by emigration relaxes competition in the former, and depletion of the resource by subsidized consumers decouples the local community in the latter.  相似文献   

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