Classical Metapopulation Research Articles

Trade-offs and Spatial Life-History Strategies in Classical Metapopulations

Trade-offs and Spatial Life-History Strategies in Classical Metapopulations

Metapopulation processes and persistence in remnant water vole populations

We examined the spatial distribution of water vole populations in four consecutive years and investigated whether the regional population processes of extinction, recolonisation and migration influence distribution and persistence. We examined how such regional processes are influenced by spatial variation in habitat quality. In addition, we assessed the relevance of metapopulation concepts for understanding the dynamics of species that deviate from classical metapopulation assumptions and developing conservation measures for them. Populations were patchy and discrete, and the patchy distribution was not static between years. Population turnover occurred even in the absence of predatory mink, which only influenced the network of populations at the end of the study. Most populations were clustered close together in the upper tributaries. Local population persistence was predominantly influenced by population size: large populations were more persistent. Recolonisation rates were influenced by isolation and habitat quality. The isolation estimates which best explained the distribution of water vole populations incorporated straight‐line distances, suggesting water voles disperse overland. The distribution of recolonised sites indicated that dispersing voles actively selected habitat on the basis of its quality. Water voles depart from some of the assumptions made by frequently used metapopulation models. In particular there is no clear binary distinction between suitable and non‐suitable habitat. Accounting for variation in habitat quality before investigating temporal changes in population distribution allowed us to demonstrate that the key metapopulation processes were important. The significance of regional population processes relative to local population processes may have increased in declining, fragmented populations compared to pristine regional populations. We hypothesise that although mink predation is likely to eventually cause regional extinction in many areas, metapopulation processes have delayed this decline. Consequently, conservation measures should take into account mink predation rates and regional population processes, before considering aspects of habitat quality.

Spatially realistic theory of metapopulation ecology.

Much of spatial ecology since the late 1960s has been dominated by two theories, the dynamic theory of island biogeography and the classical metapopulation theory. The latter theory largely replaced the former one in the 1980s, especially in conservation applications. It is only recently that ecologists have fully realized that a relatively simple general theory can be readily constructed that makes some of the simplifying assumptions of the two earlier theories unnecessary. The spatially realistic metapopulation theory thereby provides a more unified framework for spatial ecology than the island theory or the classical metapopulation theory. This article describes the application of the spatially realistic metapopulation theory to real metapopulations living in highly fragmented landscapes. I discuss the principal messages for population ecology and conservation biology, and I also place this theory into a broader context of other approaches to spatial ecology.

Inbreeding in nature: brothers and sisters, do not unite!

Inbreeding, or mating among related individuals, causes problems with viability and fertility in small captive populations. Young inbred ungulates often die in zoos, and the eggs of inbred fruit flies frequently fail to hatch in the laboratory. But what is the role of inbreeding in nature? Does it really affect patterns of population change in the wild, and could it even cause extinctions? These questions were the subject of hot debate in the 1970s. By the late 1980s, many ecologists seemed to agree that other ecological factors were more important than inbreeding in reducing population size and in extinguishing populations. However, we could now be witnessing the revival of interest in inbreeding as a factor to contend with. Whereas theoretical studies have demonstrated how the accumulation of mildly deleterious alleles can lower the fitness of small populations, experiments, both in the laboratory and in the wild, have demonstrated significant effects of inbreeding on the density of several animal and plant species.Observations in 1998 suggested that inbreeding increases the risk of population extinction in the butterfly Melitaea cinxia.1xInbreeding and extinction in a butterfly metapopulation. Saccheri, I et al. Nature. 1998; 392: 491–494Crossref | Scopus (1008)See all References1 Some of the same authors now report experimental evidence2xExperimental confirmation that inbreeding depression increases extinction risk in butterfly populations. Nieminen, M et al. Am. Nat. 2001; 157: 237–244Crossref | PubMed | Scopus (106)See all References2 that not only confirms the earlier results, but also demonstrates when and how inbreeding exerts its harmful effect. Working on small populations of Melitaea cinxia in the laboratory, Nieminen et al. found that inbreeding lowered the proportion of eggs hatched and decreased the survival of over-wintering larvae. The larvae of this butterfly spin communal winter nests, but inbreeding made them spin nests full of holes, which failed to protect them from the elements. And when Nieminen et al. introduced in- and outbred populations into the field, they found that inbred populations had a much higher risk of extinction than did outbred ones.This study suggests that inbreeding in Melitaea cinxia may reduce local survival to the point of local extinction. And as the species occurs as a set of classical metapopulations, where the extinction of small local populations is counterbalanced by the colonization of empty sites, an increased risk of local extinction might ultimately threaten the survival of the species at a landscape level.The work of Nieminen et al. is clearly a case study on a single species with many special features. In their study region, Melitaea cinxia typically occurs as very small local populations, where inbreeding between close relatives is more the rule than an exception. This should keep us from generalizing the current results too broadly. But by identifying inbreeding as a key agent in the dynamics of at least this species, Nieminen et al. do offer an eye opener. With more and more species being split into smaller and more fragmented populations, perhaps inbreeding might soon be a threat to the survival of many wild populations?

Spatial population structure in a patchily distributed beetle.

The dynamics and evolution of populations will critically depend on their spatial structure. Hence, a recent emphasis on one particular type of structure--the metapopulation concept of Levins--can only be justified by empirical assessment of spatial population structures in a wide range of organisms. This paper focuses on Aphodius fossor, a dung beetle specialized on cattle pastures. An agricultural database was used to locate nearly 50 000 local populations of A. fossor in Finland. Several independent methods were then used to quantify key processes in this vast population system. Allozyme markers and mitochondrial DNA (mtDNA) sequences were applied to examine genetic differentiation of local populations and to derive indirect estimates of gene flow. These estimates were compared to values expected on the basis of direct observations of dispersing individuals and assessments of local effective population size. Molecular markers revealed striking genetic homogeneity in A. fossor. Differentiation was only evident in mtDNA haplotype frequencies between the isolated Aland islands and the Finnish mainland. Thus, indirect estimates of gene flow agreed with direct observations that local effective population size in A. fossor is large (hundreds of individuals), and that in each generation, a substantial fraction (approximately one-fifth) of the individuals move between populations. Large local population size, extreme haplotype diversity and a high regional incidence of A. fossor all testify against recurrent population turnover. Taken together, these results provide strong evidence that the whole mainland population of A. fossor is better described as one large 'patchy population', with substantial movement between relatively persistent local populations, than as a classical metapopulation.

Distribution and abundance of dung beetles in fragmented landscapes.

Related species utilising similar resources are often assumed to show similar spatial population structures and dynamics. This paper reports substantial ecological variation within a set of Aphodius dung beetles occurring in the same patchily distributed resource, livestock dung in pastures. We show how variation in habitat and resource selectivity, in the rate of movements between pastures, and in the distribution of local population sizes all contribute to interspecific differences in spatial population structures. Local dung beetle assemblages are compared between two landscapes with different densities of pastures. In one of the landscapes, we contrast the abundances and regional distributions of Aphodius before and after 15years of rapid habitat loss. Different species show very dissimilar responses to changes in the structure of the landscape. Our results suggest that generalist Aphodius species, and specialist species with high dispersal powers, occur as large "patchy" populations in the landscape. In contrast, a strict pasture specialist species with limited dispersal powers (A. pusillus) forms classical metapopulations. At the community level, interspecific differences in spatial population structures make the local community composition a function of the structure of the surrounding landscape.

Metapopulation Dynamics and Amphibian Conservation

Abstract: In many respects, amphibian spatial dynamics resemble classical metapopulation models, in which subpopulations in breeding ponds blink in and out of existence and extinction and colonization rates are functions of pond spatial arrangement. This “ponds‐as‐patches” view of amphibian spatial dynamics is useful in several respects. First, it highlights the importance of regional and landscape processes in determining local patterns of abundance. Second, it offers a straightforward, pond‐based approach to monitoring and managing amphibian populations. For many species, however, the ponds‐as‐patches view may be an oversimplification and metapopulation structure may be more apparent than real. Changes in distribution may be caused by processes other than extinction and recolonization, and most extinctions probably result from deterministic factors, not stochastic processes. In addition, the effects of pond isolation appear to be important primarily in disturbed environments, and in many cases these isolation effects may be better explained by the distribution of terrestrial habitats than by the distribution of breeding ponds. These complications have important implications for both researchers and managers. For researchers, future efforts need to determine the mechanisms underlying patterns of abundance and distributional change and patterns in amphibian populations. For managers, effective conservation strategies must successfully balance metapopulation considerations with careful attention to local habitat quality. Furthermore, translocations and active management may be indispensable tools for conserving amphibians in landscapes containing multiple breeding ponds.

Effects of metapopulation processes on measures of genetic diversity

Many species persist as a metapopulation under a balance between the local extinction of subpopulations or demes and their recolonization through dispersal from occupied patches. Here we review the growing body of literature dealing with the genetic consequences of such population turnover. We focus our attention principally on theoretical studies of a classical metapopulation with a 'finite-island' model of population structure, rather than on 'continent-island' models or 'source-sink' models. In particular, we concern ourselves with the subset of geographically subdivided population models in which it is assumed that all demes are liable to extinction from time to time and that all demes receive immigrants. Early studies of the genetic effects of population turnover focused on population differentiation, such as measured by F(ST). A key advantage of F(ST) over absolute measures of diversity is its relative independence of the mutation process, so that different genes in the same species may be compared. Another advantage is that F(ST) will usually equilibrate more quickly following perturbations than will absolute levels of diversity. However, because F(ST) is a ratio of between-population differentiation to total diversity, the genetic effects of metapopulation processes may be difficult to interpret in terms of F(ST) on its own, so that the analysis of absolute measures of diversity in addition is likely to be informative. While population turnover may either increase or decrease F(ST), depending on the mode of colonization, recurrent extinction and recolonization is expected always to reduce levels of both within-population and species-wide diversity (piS and piT, respectively). One corollary of this is that piS cannot be used as an unbiased estimate of the scaled mutation rate, theta, as it can, with some assumptions about the migration process, in species whose demes do not fluctuate in size. The reduction of piT in response to population turnover reflects shortened mean coalescent times, although the distribution of coalescence times under extinction colonization equilibrium is not yet known. Finally, we review current understanding of the effect of metapopulation dynamics on the effective population size.

EMPIRICAL EVIDENCE FOR PREDATOR–PREY SOURCE–SINK DYNAMICS

Theory suggests that a source–sink structure for a prey species can promote the persistence of an otherwise nonpersisting predator–prey interaction. Using a heliozoan protist predator, Actinosphaerium nucleofilum, and a ciliated protozoan prey species, Tetrahymena pyriformis, we tested this prediction in laboratory microcosms. We created subdivided microcosms, each consisting of a 30-mL bottle containing predators and prey connected to a 30-mL bottle containing prey only. Prey dispersed freely through this connection, but predators did not disperse for hundreds of prey generations. The predators and prey in the subdivided microcosms persisted for over three times as long as they did in undivided 30-mL and 60-mL bottles. Our results suggest that prey rescue effects and spatial asynchrony in prey dynamics, characteristic metapopulation features, enhanced persistence in the subdivided microcosms. However, the details by which persistence was achieved closely resemble source–sink dynamics, not classic metapopulation dynamics. Evidence suggests that continuous prey immigration into predator–prey bottles from extinction-invulnerable prey-only bottles may have weakened the coupling between predator and prey dynamics and contributed to the increase in persistence. In showing that source–sink dynamics enhanced predator–prey persistence, our experiments support conclusions of metapopulation theory that point to the importance of immigration between spatially discrete populations.

Metapopulation dynamics in an aphid‐parasitoid system

AbstractMetapopulation theory makes a number of predictions concerning the effects of dispersal on the persistence of predator‐prey or host‐parasitoid systems. While the stabilising effects of dispersal have been shown in a number of laboratory studies, evidence from field studies remains scarce due to a lack of suitable model systems. I describe a host‐parasitoid system that shows a classical metapopulation structure with frequent extinctions and colonisations consisting of the aphidiid Lysiphlebus hirticornis and the aphid Metopeurum fuscoviride. Both the parasitoid and the aphid are specialists on their respective hosts. I followed the dynamics of host and parasitoid on individually marked tansy (Tanacetum vulgare) plants, the host of M. fuscoviride. Dynamics of host and parasitoid populations were characterized by frequent extinctions and colonisations. Mean longevity of aphid colonies was only 3.1 weeks. Parasitism by L. hirticornis was a main cause of extinction for the aphid as rates of parasitism often reached 100%, in particular towards the end of the field season. Patchiness in this system occurs at two spatial scales. Aphid colonies form on single tansy ramets = shoots but movements of aphid individuals among ramets within a particular tansy genet are frequent. Because aphids can persist on a genet for a large numer of generations, it is argued that local populations form on genets rather than ramets. The number of host and parasitoid extinctions described in this study exceeds the number of extinctions usually observed in field studies of host‐parasitoid metapopulations. It is suggested that aphid‐parasitoid systems such as the one studied in this paper may be good models to test the predictions of metapopulation theory.

The influence of patch demographics on metapopulations, with particular reference to successional landscapes

The classical view of metapopulations relates the regional abundance of a species to the balance between the extinction and colonization dynamics of identical local populations. Species in successional landscapes may represent the most appropriate examples of classical metapopulations. However, Levins‐type metapopulation models do not explicitly separate population loss due to successional habitat change from other causes of extinction. A further complication is that the chance of population loss due to successional habitat change may be related to the age of a patch. I developed simple patch occupancy models to include succession and included consideration of patch age structure to address two related questions: what are the implications of changes in patch demographic rates and when is a move to a structured patch occupancy model justified? Age‐related variation in patch demography could increase or decrease the equilibrium fraction of the available habitat occupied by a species when compared to the predictions of an unstructured model. Metapopulation persistence was enhanced when the age class of patches with the highest species occupancy suffered relatively low losses to habitat succession. Conversely, when the age class of patches with the highest species occupancy also had relatively high successional loss rates, extinction thresholds were higher that would be predicted by a simple unstructured model. Hence age‐related variation in patch successional rate introduces biases into the predictions of simple unstructured models. Such biases can be detected from field surveys of the fraction of occupied and unoccupied patches in each age class. Where a bias is demonstrated, unstructured models will not be adequate for making predictions about the effects of changing parameters on metapopulation size. Thinking in successional terms emphasizes how landscapes might be managed to enhance or reduce the patch occupancy by any particular metapopulation

Immigration and the Maintenance of Local Species Diversity.

Explaining the maintenance of high local species diversity in communities governed by competition for space has been a long-standing problem in ecology. We present a simple theoretical model to explore the influence of immigration from an external source on local coexistence, species abundance patterns, and ecosystem processes in plant communities. The model is built after classical metapopulation models but is applied to competition for space between individuals and includes immigration by a propagule rain and an extinction threshold for rare species. Our model shows that immigration can have a huge effect on local species diversity in competitive communities where competition for space would lead to the exclusion of all but one species if the community were closed. Local species richness is expected to increase strongly when immigration intensity increases beyond the threshold required for the successful establishment of one or a few individuals. Community structure and species relative abundances are also expected to change markedly with immigration intensity. Increasing immigration causes total space occupation by the community to increase but primary productivity on average to either decrease or stay constant with increasing diversity, depending on the relation between immigration and local reproduction rates. These results stress the need for a regional perspective to understand the processes that determine species diversity, species abundance patterns, and ecosystem functioning in local communities.

HYDROTHERMAL-VENT ALVINELLID POLYCHAETE DISPERSAL IN THE EASTERN PACIFIC. 2. A METAPOPULATION MODEL BASED ON HABITAT SHIFTS.

Marine organisms typically fall into two main categories: those with a high level of population structuring and those with a low one. The first are often found to be poor dispersers, following isolation by distance or stepping-stone theoretical predictions. The second are commonly associated with high-dispersal taxa and are best described by the island model. Deep-sea hydrothermal vent systems represent a good model for studying one-dimensional metapopulations. Whereas isolation by distance might be expected to be the rule in such a system for species with limited dispersal capabilities, a biological paradox can be observed: an apparent genetic homogeneity in some vent species with short-scale dispersal potential, in a one-dimensional fragmented habitat. This can be explained if one key assumption of the existing models is not met: gene flow between populations and genetic drift may not have the time to equilibrate. Geophysical models revealed that hydrothermal convection is intrinsically unstable, inducing processes of coalescence or splitting of venting areas in a chaotic manner. This is likely to generate frequent extinctions and recolonizations. Theoretical genetic predictions derived from extinctions/recolonizations cannot satisfactorily model a situation where habitat shifts are frequent and constantly affect the metapopulation equilibrium. Because neither the island and the stepping-stone models nor the classical metapopulation models resemble the hydrothermal vent reality, we present here a realistic model developed to provide a compromise between existing conceptual models and what is currently known of the biology and ecology of one of the most peculiar and best-studied vent species, the polychaete Alvinella pompejana. This model allows us to define the boundaries between which the metapopulation is evolutionary stable in an unstable context. Simulations show different patterns in which metapopulation size and recolonization vary but reach an equilibrium despite chaotic vent extinctions. In contrast, the model also shows that displacing habitat continuously affects the equilibrium between gene flow and drift. As a consequence, the time required to balance these evolutionary forces can never be attained, leading to chaotic fluctuations in F-statistics. Those fluctuations are mainly due to stochastic changes of the interpatch distance which affect migration rates. The shifting of active zones of venting can episodically counterbalance differentiation and allow a long-term genetic homogenization at the ridge scale. These findings lead to a new concept in which the exchanges between populations would mainly depend on the habitat's movements along the ridge axis rather than the organim's dispersal. We therefore propose a new model based on patch-network displacements in which transient contact zones allow low levels of gene flow throughout the metapopulation.

Spatially structured populations: how much do they match the classic metapopulation concept?

In the classic metapopulation concept a specific range of animal movements is assumed, not too large and not too small. Thus, knowledge of animal mobility is necessary to determine the degree to which a given population matches a specific metapopulation model. It seems that usually small mammal mobility is underestimated, and this has important consequences for the way we view metapopulation dynamics. Data on small mammal movements (Clethrionomys glareolus and Apodemus flavicollis) are presented in this paper. The material was collected in a two year study in western Poland using a set of six woodlots of different sizes and degree of isolation, located among agricultural fields. Various methods were used in the study: colored bait, live-trapping, and radio-telemetry. It is suggested that the populations under study match the concept of patchy population (sensu Harrison 1991), being poorly isolated in individual patches and with the range of animal movements encompassing the whole set of patches. Moreover, the use of patches changes between seasons according to changing needs and/or resource abundance in the woodlots. Density and composition of local populations may be influenced both by the patch area and its isolation, and also by the filtering effect of the matrix that depends on the season. In this context it is pointed out that more attention should be paid to the matrix, both in research practice and conservation as it is a factor influencing population functioning in quantitative and qualitative ways. It is suggested that any generalizations about population spatial organization may be impossible without more detailed knowledge of long distance movements of focal animals and their use of matrix.

Immigration and the Maintenance of Local Species Diversity

Explaining the maintenance of high local species diversity in communities governed by competition for space has been a long‐standing problem in ecology. We present a simple theoretical model to explore the influence of immigration from an external source on local coexistence, species abundance patterns, and ecosystem processes in plant communities. The model is built after classical metapopulation models but is applied to competition for space between individuals and includes immigration by a propagule rain and an extinction threshold for rare species. Our model shows that immigration can have a huge effect on local species diversity in competitive communities where competition for space would lead to the exclusion of all but one species if the community were closed. Local species richness is expected to increase strongly when immigration intensity increases beyond the threshold required for the successful establishment of one or a few individuals. Community structure and species relative abundances are also expected to change markedly with immigration intensity. Increasing immigration causes total space occupation by the community to increase but primary productivity on average to either decrease or stay constant with increasing diversity, depending on the relation between immigration and local reproduction rates. These results stress the need for a regional perspective to understand the processes that determine species diversity, species abundance patterns, and ecosystem functioning in local communities.

Genetic variation in a freshwater bryozoan. I: Populations in the Thames basin, UK.

In spite of increasing interest in metapopulation dynamics, the genetic consequences of a metapopulation structure remain poorly understood. Here we examine the metapopulation genetic structure of the colonial, facultatively sexual freshwater bryozoan Cristatella mucedo, in the Thames basin of southern England, UK. Populations from nine sites were sampled and colonies genetically characterized using random amplified polymorphic DNA (RAPD)-PCR. A total of 78 different clones was detected over all sites. Despite the large number of clones, genetic distances among clones both within and among sites were very small. Nonetheless, no clone was present at more than one site even though C. mucedo has an asexual dispersal propagule, and clones strongly clustered by sites. No consistent pattern of clonal structure was evident, with both the number and equitability of clones varying greatly among sites. Although sites were genetically distinct, population genetic regions were absent, and a Mantel test indicated that there was no relationship between geographical distances among sites and genetic distances among populations. Our results indicate that C. mucedo exists as a classical metapopulation in the Thames basin, with dispersal independent of distance and all sites contributing to the genetic diversity of the metapopulation.

Connecting the Parameters of Local Extinction and Metapopulation dynamics

This paper explores the correspondence between the parameters of an extinction model analysed by Lande, Foley and Middleton et al. and the parameters of the incidence function model of metapopulation dynamics. The parameters of the extinction model, the intrinsic rate of population increase (r), its variance (v) and the population ceiling (K), can be mapped to the parameters of the incidence function model describing the scaling of the probability of local extinction (E) with patch area (A), E = eA X, via the equations s = x and r = eDs/s, where s = 2r/v, D is population density and K= DA. I explore this correspondence with two empirical examples, a mainlandisland metapopulation of the European common shrew (Sorex araneus) on islands in lakes and a classical metapopulation of the American pika (Ochotona princess. The most robust result is the correspondence x = 2r/v, which value decreases with increasing strength of environmental stochasticity. Thus the impact of environmental stochasticity on population dynamics can, in principle, be inferred from the pattern of habitat patch occupancy in a metapopulation.

Long‐Term Dynamics in a Metapopulation of the American Pika

A 20-yr study of a metapopulation of the American pika revealed a regional decline in occupancy in one part of a large network of habitat patches. We analyze the possible causes of this decline using a spatially realistic metapopulation model, the incidence function model. The pika metapopulation is the best-known mammalian example of a classical metapopulation with significant population turnover, and it satisfies closely the assumptions of the incidence function model, which was parameterized with data on patch occupancy. The model-predicted incidences of patch occupancy are consistent with observed incidences, and the model predicts well the observed turnover rate between four metapopulation censuses. According to model predictions, the part of the metapopulation where the decline has been observed is relatively unstable and prone to large oscillations in patch occupancy, whereas the other part of the metapopulation is predicted to be persistent. These results demonstrate how extinction-colonization dynamics may produce spatially correlated patterns of patch occupancy without any spatially correlated processes in local dynamics or extinction rate. The unstable part of the metapopulation gives an empirical example of multiple quasi equilibria in metapopulation dynamics. Phenomena similar to those observed here may cause fluctuations in species' range limits.

Pattern Formation in a Patch Occupancy Metapopulation Model: a Cellular Automata Approach

The explicit consideration of space in ecological research is of paramount importance to understand the structure and functioning of ecological systems. In this paper we develop a simple spatially explicit metapopulation model in which colonization is constant and independent of the number of occupied patches (i.e. propagule-rain effect, Gotelli, 1991). Extinction, on the other hand, is modelled as a stochastic process whose intensity depends on the number of occupied patches in the neighborhood of each focal patch. Our model is the CA counterpart of two classical patch occupancy metapopulation models. We analytically prove this by showing that our CA converges to the differential equation in the mean-field approximation. The asymptotic behaviour of the system, expressed as the proportion of occupied patches, agrees with the equilibrium proportion of patches derived by using ODEs. In both models, the existence of a rescue-effect increases the range of extinction and colonization parameters over which the system attains complete occupancy of patches. However, in our model this result is strongly influenced by the degree of coupling among patches and is apparent only for local interactions. With local interactions and particular parameter values of colonization and extinction, self-organized spatio-temporal patterns emerge with a fractal-like clustering, even though the environment is spatially homogeneous. Our results point out that the importance of being spatial and discrete (Durrett & Levin, 1994a) in our model is a result of local interactions.

Long-Term Dynamics in a Metapopulation of the American Pika

A 20‐yr study of a metapopulation of the American pika revealed a regional decline in occupancy in one part of a large network of habitat patches. We analyze the possible causes of this decline using a spatially realistic metapopulation model, the incidence function model. The pika metapopulation is the best‐known mammalian example of a classical metapopulation with significant population turnover, and it satisfies closely the assumptions of the incidence function model, which was parameterized with data on patch occupancy. The model‐predicted incidences of patch occupancy are consistent with observed incidences, and the model predicts well the observed turnover rate between four metapopulation censuses. According to model predictions, the part of the metapopulation where the decline has been observed is relatively unstable and prone to large oscillations in patch occupancy, whereas the other part of the metapopulation is predicted to be persistent. These results demonstrate how extinction‐colonization dynamics may produce spatially correlated patterns of patch occupancy without any spatially correlated processes in local dynamics or extinction rate. The unstable part of the metapopulation gives an empirical example of multiple quasi equilibria in metapopulation dynamics. Phenomena similar to those observed here may cause fluctuations in species' range limits.