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.     

Colonization and extinction dynamics of an annual plant metapopulation in an urban environment

The metapopulation framework considers that the spatiotemporal distribution of organisms results from a balance between the colonization and extinction of populations in a suitable and discrete habitat network. Recent spatially realistic metapopulation models have allowed patch dynamics to be investigated in natural populations but such models have rarely been applied to plants. Using a simple urban fragmented population system in which favourable habitat can be easily mapped, we studied patch dynamics in the annual plant Crepis sancta (Asteraceae). Using stochastic patch occupancy models (SPOMs) and multi‐year occupancy data we dissected extinction and colonization patterns in our system. Overall, our data were consistent with two distinct metapopulation scenarios. A metapopulation (sensu stricto) dynamic in which colonization occurs over a short distance and extinction is lowered by nearby occupied patches (rescue effect) was found in a set of patches close to the city centre, while a propagule rain model in which colonization occurs from a large external population was most consistent with data from other networks. Overall, the study highlights the importance of external seed sources in urban patch dynamics. Our analysis emphasizes the fact that plant distributions are governed not only by habitat properties but also by the intrinsic properties of colonization and dispersal of species. The metapopulation approach provides a valuable tool for understanding how colonization and extinction shape occupancy patterns in highly fragmented plant populations. Finally, this study points to the potential utility of more complex plant metapopulation models than traditionally used for analysing ecological and evolutionary processes in natural metapopulations.     

Modelling the spatial dynamics and persistence of the leaf beetle Gonioctena olivacea in dynamic habitats

In dynamic landscapes natural and anthropogenic disturbance as well as succession are responsible for the emergence and subsequent disappearance of suitable habitat patches. Species inhabiting such landscapes are faced with varying number and spatial configuration of patches. A stochastic, spatially explicit simulation model was developed in order to analyse the persistence of the leaf beetle Gonioctena olivacea in a system of dynamic patches of its host plant Cytisus scoparius . The model was parameterized with data from a three-year field study on the spatial configuration, distribution, and turnover of the host plant patches as well as the patch occupancy, extinction, and colonization rates of the beetle. The simulations showed large fluctuations in the occurrence of the beetle in the patches. High levels of occupancy were related to high aggregation of the patches within the landscape. The velocity of patch turnover was found to have a severe effect on the persistence of the beetle metapopulation. Enhancing the turnover rate by only a few patches, the mean time to extinction decreases rapidly. Moreover, the results revealed that not necessarily an effect of connectivity can be detected in the analysis of occupancy patterns in dynamic landscapes, although the colonization of patches is clearly connectivity-dependent. In general, this modelling study demonstrates the importance of detailed information on patch turnover. The amount and spatial distribution of suitable habitat is a major driver of metapopulation dynamics of species in dynamic landscapes.     

The effective size of a metapopulation living in a heterogeneous patch network

I analyze stochastic patch occupancy models (SPOMs), which record habitat patches as empty or occupied. A problem with SPOMs has been that if the spatial structure of a heterogeneous habitat patch network is taken into account, the computational effort needed to analyze a SPOM grows as a power of 2n, where n is the number of habitat patches. I propose a computationally feasible approximation method, which approximates the behavior of a heterogeneous SPOM by an "ideal" metapopulation inhabiting a network of identical and equally connected habitat patches. The transformation to the ideal metapopulation is based on weighting the individual patch occupancies by the dynamic values of the habitat patches, which may be calculated from the deterministic mean-field approximation of the original SPOM. Conceptually, the method resembles the calculation of the effective size of a population in the context of population genetics. I demonstrate how the method may be applied to SPOMs with flexible structural assumptions and with spatially correlated and temporally varying parameter values. I apply the method to a real habitat patch network inhabited by the Glanville fritillary butterfly, illustrating that the metapopulation dynamics of this species are essentially driven by temporal variability in the environmental conditions.     

Testing metapopulation concepts: effects of patch characteristics and neighborhood occupancy on the dynamics of an endangered lagomorph

Metapopulation ecology is a field that is richer in theory than in empirical results. Many existing empirical studies use an incidence function approach based on spatial patterns and key assumptions about extinction and colonization rates. Here we recast these assumptions as hypotheses to be tested using 18 years of historic detection survey data combined with four years of data from a new monitoring program for the Lower Keys marsh rabbit. We developed a new model to estimate probabilities of local extinction and colonization in the presence of nondetection, while accounting for estimated occupancy levels of neighboring patches. We used model selection to identify important drivers of population turnover and estimate the effective neighborhood size for this system. Several key relationships related to patch size and isolation that are often assumed in metapopulation models were supported: patch size was negatively related to the probability of extinction and positively related to colonization, and estimated occupancy of neighboring patches was positively related to colonization and negatively related to extinction probabilities. This latter relationship suggested the existence of rescue effects. In our study system, we inferred that coastal patches experienced higher probabilities of extinction and colonization than interior patches. Interior patches exhibited higher occupancy probabilities and may serve as refugia, permitting colonization of coastal patches following disturbances such as hurricanes and storm surges. Our modeling approach should be useful for incorporating neighbor occupancy into future metapopulation analyses and in dealing with other historic occupancy surveys that may not include the recommended levels of sampling replication.     

Metapopulation theory for fragmented landscapes

We review recent developments in spatially realistic metapopulation theory, which leads to quantitative models of the dynamics of species inhabiting highly fragmented landscapes. Our emphasis is in stochastic patch occupancy models, which describe the presence or absence of the focal species in habitat patches. We discuss a number of ecologically important quantities that can be derived from the full stochastic models and their deterministic approximations, with a particular aim of characterizing the respective roles of the structure of the landscape and the properties of the species. These quantities include the threshold condition for persistence, the contributions that individual habitat patches make to metapopulation dynamics and persistence, the time to metapopulation extinction, and the effective size of a metapopulation living in a heterogeneous patch network.     

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     

Spatial and temporal variation in patch occupancy and population density in a model system of an arctic Collembola species assemblage

We employed an experimental model system to investigate the mechanisms underlying patterns of patch occupancy and population density in a high arctic assemblage of Collembola species inhabiting a sedge tussock landscape on Svalbard. The replicate model systems consisted of 5 cores of the tussocks (habitat patches) imbedded in a barren matrix. Four of the patches were open so that animals could migrate between them, while there was one closed patch per system to test the effect of migration on extinction rate. There were model systems of two types: one with long and one with short inter‐patch distances to test the effect of patch isolation on colonisation and extinction rates. Each of the four most common collembolan species at the field site were introduced to two open patches per system (source patches), with the other two functioning as colonisation patches for the species. The experiment was run in an ecotrone over three identical, simulated arctic summers separated by winters of 3 weeks. Six replicates of systems with short and long inter‐patch distances were sampled at the end of each summer. The species varied markedly in their performance in both open arenas and closed patches, indicating differential responses to patch humidity, consistent with their differential distribution along the moisture gradient in the field site. The extinction – colonisation dynamics differed markedly between species as predicted from our field studies. This could partly be ascribed to differential dispersal and colonisation ability, but also to different tolerance to spatially variable patch quality and/or tendency for aggregative behaviour. Three of the species exhibited dynamics that superficially resemble what could be expected from classical metapopulation dynamics. However, there was a striking discrepancy between what would be expected from the effect of migration on the extinction rate of isolated patches (in particular closed patches) and the observed rates. Thus, metapopulation processes, such as stochastic colonisation and extinction events due to demographic stochasticity, were relatively unimportant compared to other sources of spatial variability among which subtle differences in patch quality are probably most important. We discuss the value of combining field studies with model system experiments, in particular when habitat quality cannot easily be measured in the field. However, our field and laboratory studies also emphasise the need for a thorough knowledge of species‐specific life history traits for making biologically sound interpretations based on both observational and experimental data.     

用空间直观模型是否足以从斑块占据性资料中推断集合种群的动态过程?

在集合种群的研究中,经常要根据空间占据性数据应用斑块模型来推断种群的动态过程,在保护生物学应用中,斑块占据性模型的参数估测对于阐释集合种群动态和预测种群对生境破坏的反应极为重要。我们探讨了一种广泛应用的空间直观模型——率函数模型(Incidence function model)中参数估测的不确定性问题,通过构建由50个斑块组成的网络和两个假想的已知参数的集合种群,应用模拟模型产生集合种群随时间变化的斑块占据性数据系列：即快照(snapshot)。然后,根据这些快照,应用率函数模型和最大似然法估测种群动态参数。此外,我们还给出了传统的率函数模型的一个变形,这个变形包含了目标区效应(Target area effect)：即一个斑块的占据概率不但取决于空间隔离度,也取决于斑块本身面积的大小。结果表明：根据同一个集合种群不同的快照所估测的参数可以有很大差异,一个快照得出的参数提示的是占据性强但存活率低的集合种群,而另一个快照可能反映的是一个占据性弱但存活率高的集合种群。应用传统的率函数模型于一个包含了目标区效应的集合种群,导致斑块大小相关的灭绝率参数估测的正偏差。因此,仅根据一个快照的空间占据性数据来推测集合种群的过程有很大的不确定性[动物学报49(6)：787～794,2003]。     

Effect of habitat fragmentation on dispersal in the butterfly Proclossiana eunomia

Comparison of dispersal rates of the bog fritillary butterfly between continuous and fragmented landscapes indicates that between patch dispersal is significantly lower in the fragmented landscape, while population densities are of the same order of magnitude. Analyses of the dynamics of the suitable habitat for the butterfly in the fragmented landscape reveal a severe, non linear increase in spatial isolation of patches over a time period of 30 years (i.e. 30 butterfly generations), but simulations of the butterfly metapopulation dynamics using a structured population model show that the lower dispersal rates in the fragmented landscape are far above the critical threshold leading to metapopulation extinction. These results indicate that changes in individual behaviour leading to the decrease of dispersal rates in the fragmented landscape were rapidly selected for when patch spatial isolation increased. The evidence of such an adaptive answer to habitat fragmentation suggests that dispersal mortality is a key factor for metapopulation persistence in fragmented landscapes. We emphasise that landscape spatial configuration and patch isolation have to be taken into account in the debate about large-scale conservation strategies.     

Population turnover and habitat dynamics in Notonecta (Hemiptera: Notonectidae) metapopulations

Simple metapopulation models assume that local populations occur in patches of uniform quality habitat separated by non-habitat. However field metapopulations tend to show considerable spatial and temporal variation in patch quality, and hence probability of occupancy. This may have implications for the adequacy of simple metapopulation models in describing and predicting regional population dynamics of natural systems. This study investigated the effects of habitat characteristics on landscape-scale occupancy dynamics of two species of backswimmer (Notonecta, Hemiptera: Notonectidae) in small freshwater ponds. The results demonstrated clear links between habitat, pond occupancy and population turnover, particularly local extinction. There were considerable changes in the habitat of individual ponds between years, but local changes were not spatially correlated and the frequency distribution of habitat conditions at the landscape level remained similar in different years. Stable occupancy levels of Notonecta species appears to result from a balance of the rates of creation and loss of suitable habitat due to spatially uncorrelated habitat change. Systems such as this, where turnover is driven by habitat dynamics, demonstrate the potential value of incorporating the dynamics of habitat change into metapopulation models. Such developments are likely to improve predictions of landscape-scale occupancy dynamics, whilst also allowing patch-level predictions of occupancy, based on local habitat conditions. Received: 18 August 1999 / Accepted: 3 December 1999     

Single-species metapopulation dynamics: concepts, models and observations

This paper outlines a conceptual and theoretical framework for single-species metapopulation dynamics based on the Levins model and its variants. The significance of the following factors to metapopulation dynamics are explored: evolutionary changes in colonization ability; habitat patch size and isolation; compensatory effects between colonization and extinction rates; the effect of immigration on local dynamics (the rescue effect); and heterogeneity among habitat patches. The rescue effect may lead to alternative stable equilibria in metapopulation dynamics. Heterogeneity among habitat patches may give rise to a bimodal equilibrium distribution of the fraction of patches occupied in an assemblage of species (the core-satellite distribution). A new model of incidence functions is described, which allows one to estimate species' colonization and extinction rates on islands colonized from mainland. Four distinct kinds of stochasticity affecting metapopulation dynamics are discussed with examples. The concluding section describes four possible scenarios of metapopulation extinction.     

Quantitative predictions for patch occupancy of capercaillie in fragmented habitats

A major conclusion of studying metapopulation biology is that species conservation should favor regional rather than local population persistence. Regional persistence is tightly linked to size, spatial configuration and quality of habitat patches. Hence it is important for the management of endangered species that priority patches can be identified. We developed a predictive model of patch occupancy by capercaillie, a threatened grouse species, based on a single snapshot of data. We used logistic regression to predict patch occupancy as a function of patch size, isolation, connectivity, relative altitude, and biogeographical area. The probability of a patch being occupied increased with patch size and increasing altitude, and decreased with increasing distance to the next occupied patch. Patch size was the most important predictor although occupied patches varied considerably in size. Our model only uses data on the number, size and spatial configuration of habitat patches. It is a useful tool to designate priority areas for conservation, i.e. large core patches with high resilience in habitat quality, smaller island‐patches that still have high probability of being inhabited or becoming recolonised, and patches functioning as “stepping stones”. If capercaillie is to be preserved, habitat suitability needs to be maintained in a functional network of patches that account for size and inter‐patch distance thresholds as found in this study. We suggest that similar area‐isolation relationships are valid for almost any region within the distribution range of capercaillie. The thresholds for occupancy are however likely to depend on characteristics of the respective landscape. The outcome of our study emphasises the need for future investigations that explore the relationship between patch occupancy, matrix quality and its resistance to dispersing individuals.     

Colonization–extinction dynamics of epixylic lichens along a decay gradient in a dynamic landscape

Metapopulation models are often used for understanding and predicting species dynamics in fragmented landscapes. Several models have been proposed depending on e.g. the relative importance of patch dynamics on the metapopulation dynamics. Dead wood is a dynamic substrate patch, and species that are confined to such patches have experienced a high degree of habitat loss in managed forests. Little is, however, known about how the population dynamics of epixylic species are affected by the fast dynamics of their substrate patches. We quantified the effect of local patch conditions and metapopulation processes on colonizations and extinctions of epixylic lichen species in a managed boreal forest landscape. This was done by twice surveying seven lichen metapopulations on 293 stumps in 30 stands of ages covering the duration of the dynamic patches (stumps). We also investigated the relative importance of local stochastic extinctions from stumps that remained available, and deterministic extinctions due to stump surface disappearance. We found importance of a decay gradient, surrounding metapopulation size, and local population sizes, in driving the colonization–extinction dynamics of epixylic lichens. The species were sorted along the stump decay gradient. Increasing surrounding metapopulation size was associated with increased colonization rates, and increasing local population size decreased lichen extinction rates. Finally, both local stochastic extinctions and deterministic extinctions due to patch disappearance occur, confirming that the long‐term persistence of epixylic lichens depends on colonization rates that compensate for stochastic population extinctions as well as deterministic extinctions.     

Do invasive species undergo metapopulation dynamics? A case study of the invasive Caspian gull,Larus cachinnans,in Poland

Aim The mechanisms of initial dispersal and habitat occupancy by invasive alien species are fundamental ecological problems. Most tests of metapopulation theory are performed on local population systems that are stable or in decline. In the current study we were interested in the usefulness of metapopulation theory to study patch occupancy, local colonization, extinction and the abundance of the invasive Caspian gull (Larus cachinnans) in its initial invasion stages. Location Waterbodies in Poland. Methods Characteristics of the habitat patches (waterbodies, 35 in total) occupied by breeding pairs of Caspian gulls and an equal sample of randomly selected unoccupied patches were compared with t‐tests. Based on presence–absence data from 1989 to 2006 we analysed factors affecting the probability of local colonization, extinction and the size of local populations using generalized linear models. Results Occupied habitat patches were significantly larger and less isolated (from other habitat patches and other local populations) and were located closer to rivers than empty patches. The proximity of local food resources (fish ponds, refuse dumps) positively affected the occurrence of breeding pairs. The probability of colonization was positively affected by patch area, and negatively by distances to fish ponds, nearest habitat patch, nearest breeding colony and to a river, and by higher forest cover around the patch boundaries. The probability of extinction was lower in patches with a higher number of breeding pairs and with a greater area of islets. The extinction probability increased with distances to other local populations, other habitat patches, fish ponds and to refuse dumps and with a higher cover of forest around the patch boundaries. The size of the local population decreased with distances to the nearest habitat patch, local population, river, fish pond and refuse dump. Local abundance was also positively affected by the area of islets in the patch. Main conclusions During the initial stages of the invasion of Caspian gulls in Poland the species underwent metapopulation‐like dynamics with frequent extinctions from colonized habitat patches. The results prove that metapopulation theory may be a useful conceptual framework for predicting which habitats are more vulnerable to invasion.     

Patch occupancy of North American mammals: is patchiness in the eye of the beholder?

Aim Intraspecific variation in patch occupancy often is related to physical features of a landscape, such as the amount and distribution of habitat. However, communities occupying patchy environments typically exhibit non‐random distributions in which local assemblages of species‐poor patches are nested subsets of assemblages occupying more species‐rich patches. Nestedness of local communities implies interspecific differences in sensitivity to patchiness. Several hypotheses have been proposed to explain interspecific variation in responses to patchiness within a community, including differences in (1) colonization ability, (2) extinction proneness, (3) tolerance to disturbance, (4) sociality and (5) level of adaptation to prevailing environmental conditions. We used data on North American mammals to compare the performance of these ‘ecological’ hypotheses and the ‘physical landscape’ hypothesis. We then compared the best of these models against models that scaled landscape structure to ecologically relevant attributes of individual species. Location North America. Methods We analysed data on prevalence (i.e. proportion of patches occupied in a network of patches) and occupancy for 137 species of non‐volant mammals and twenty networks consisting of four to seventy‐five patches. Insular and terrestrial networks exhibited significantly different mean levels of prevalence and occupancy and thus were analysed separately. Indicator variables at ordinal and family levels were included in models to correct for effects caused by phylogeny. Akaike's information criterion was used in conjunction with ordinary least squares and logistic regression to compare hypotheses. Results A patch network's physical structure, indexed using patch area and isolation, received the greatest support among models predicting the prevalence of species on insular networks. Niche breadth (diet and habitat) received the greatest support for predicting prevalence of species occupying terrestrial networks. For both insular and terrestrial systems, physical features (patch area and isolation) received greater support than any of the ecological hypotheses for predicting species occupancy of individual patches. For terrestrial systems, scaling patch area by its suitability to a focal species and by individual area requirements of the species, and scaling patch isolation by species‐specific dispersal ability and niche breadth, resulted in models of patch occupancy that were superior to models relying solely on physical landscape features. For all selected models, unexplained levels of variation were high. Main conclusions Stochasticity dominated the systems we studied, indicating that random events are probably quite important in shaping local communities. With respect to deterministic factors, our results suggest that forces affecting species prevalence and occupancy may differ between insular and terrestrial systems. Physical features of insular systems appeared to swamp ecological differences among species in determining prevalence and occupancy, whereas species with broad niches were disproportionately represented in terrestrial networks. We hypothesize that differential extinction over long time periods in highly variable networks has driven nestedness of mammalian communities on islands, whereas differential colonization over shorter time‐scales in more homogeneous networks probably governed the local structure of terrestrial communities. Our results also demonstrate that integration of a species' ecological traits with physical features of a patch network is superior to reliance on either factor separately when attempting to predict the species' probability of patch occupancy in terrestrial systems.     

Spatially structured metapopulation models: global and local assessment of metapopulation capacity

We model metapopulation dynamics in finite networks of discrete habitat patches with given areas and spatial locations. We define and analyze two simple and ecologically intuitive measures of the capacity of the habitat patch network to support a viable metapopulation. Metapopulation persistence capacity lambda(M) defines the threshold condition for long-term metapopulation persistence as lambda(M)>delta, where delta is defined by the extinction and colonization rate parameters of the focal species. Metapopulation invasion capacity lambda(I) sets the condition for successful invasion of an empty network from one small local population as lambda(I)>delta. The metapopulation capacities lambda(M) and lambda(I) are defined as the leading eigenvalue or a comparable quantity of an appropriate "landscape" matrix. Based on these definitions, we present a classification of a very general class of deterministic, continuous-time and discrete-time metapopulation models. Two specific models are analyzed in greater detail: a spatially realistic version of the continuous-time Levins model and the discrete-time incidence function model with propagule size-dependent colonization rate and a rescue effect. In both models we assume that the extinction rate increases with decreasing patch area and that the colonization rate increases with patch connectivity. In the spatially realistic Levins model, the two types of metapopulation capacities coincide, whereas the incidence function model possesses a strong Allee effect characterized by lambda(I)=0. For these two models, we show that the metapopulation capacities can be considered as simple sums of contributions from individual habitat patches, given by the elements of the leading eigenvector or comparable quantities. We may therefore assess the significance of particular habitat patches, including new patches that might be added to the network, for the metapopulation capacities of the network as a whole. We derive useful approximations for both the threshold conditions and the equilibrium states in the two models. The metapopulation capacities and the measures of the dynamic significance of particular patches can be calculated for real patch networks for applications in metapopulation ecology, landscape ecology, and conservation biology.     

Host–parasitoid dynamics in a fragmented landscape: Holly trees,holly leaf miners and their parasitoids

Many species inhabit fragmented landscapes, where units of resource have a patchy spatial distribution. While numerous studies have investigated how the incidence and dynamics of individual species are affected by the spatial configuration and landscape context of habitat patches, fewer studies have investigated the dynamics of multiple interacting resource and consumer species in patchy landscapes. We describe a model system for investigating host–parasitoid dynamics in a patchy landscape: a network of 166 holly trees, a specialised herbivore of holly (the leaf miner, Phytomyza ilicis (Curtis, 1948)), and its suite of parasitoids. We documented patch occupancy by P. ilicis, its density within patches, and levels of parasitism over a 6-year period, and manipulated patch occupancy by creating artificially vacant habitat patches. Essentially all patches were occupied by the herbivore in each year, suggesting that metapopulation dynamics are unlikely to occur in this system. The main determinants of densities for P. ilicis and its parasitoids were resource availability (patch size and host density, respectively). While P. ilicis is apparently not restricted by the spatial distribution of resources, densities of its parasitoids showed a weaker positive relationship with host density in more isolated patches. In patches where local extinctions were generated experimentally, P. ilicis densities and levels of parasitism recovered to pre-manipulation levels within a single generation. Furthermore, patch isolation did not significantly affect re-colonisation by hosts or parasitoids. Analysing the data at a variety of spatial scales indicates that the balance between local demography and dispersal may vary depending on the scale at which patches are defined. Taken together, our results suggest that the host and its parasitoids have dispersal abilities that exceed typical inter-patch distances. Patch dynamics are thus largely governed by dispersal rather than within-patch demography, although the role of demography is higher in larger patches.     

Spatial heterogeneity, source-sink dynamics, and the local coexistence of competing species

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.