Dispersal and establishment both limit colonization during primary succession on a glacier foreland

Plant colonization can be limited by lack of seeds or by factors that reduce establishment. The role of seed limitation in community assembly is being increasingly recognized, but in early primary succession, establishment failure is still considered more important. We studied the factors limiting colonization on the foreland of Coleman Glacier, Washington, USA, to determine the importance of seed and establishment limitation during primary succession. We also evaluated the effects of seed predation, drought, and existing vegetation on establishment. We planted seeds of seven species into plots of four different ages and found evidence that both seed and establishment limitation are strong in early succession. We also found that seed and establishment limitation both remained high in later stages of succession. Seed predation reduced establishment for most species and some evidence suggested that drought and existing vegetation also limit establishment. Because both dispersal and establishment failure restrict colonization in recently exposed habitat, late-seral forest species may have a difficult time migrating upward in response to global climate change. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Disturbance,coral reef communities,and changing ecological paradigms

We examine changing ecological theory regarding the role of disturbance in natural communities and relate past and emerging paradigms to coral reefs. We explore the elements of this theory, including patterns (diversity, distribution, and abundance) and processes (competition, succession, and disturbance), using currently evolving notions concerning matters of scale (temporal and spatial), local versus regional species richness, and the equilibrium versus nonequilibrium controversy. We conclude that any attempt to categorize coral reef communities with respect to disturbance regimes will depend on the question being asked and the desired level of resolution: local assemblage versus regional species pool, successional versus geological time, and on the taxonomic and tropic affinities of species included in the study. As with many communities in nature, coral reefs will prove to be mosaics of species assemblages with equilibrial and nonequilibrial dynamics.     

Plant interspecific differences in arbuscular mycorrhizal colonization as a result of soil carbon addition

Soil nutrient availability and colonization by arbuscular mycorrhizal fungi are important and potentially interacting factors shaping vegetation composition and succession. We investigated the effect of carbon (C) addition, aimed at reducing soil nutrient availability, on arbuscular mycorrhizal colonization. Seedlings of 27 plant species with different sets of life-history traits (functional group affiliation, life history strategy and nitrophilic status) were grown in pots filled with soil from a nutrient-rich set-aside field and amended with different amounts of C. Mycorrhizal colonization was progressively reduced along the gradient of increasing C addition in 17 out of 27 species, but not in the remaining species. Grasses had lower colonization levels than forbs and legumes and the decline in AM fungal colonization was more pronounced in legumes than in other forbs and grasses. Mycorrhizal colonization did not differ between annual and perennial species, but decreased more rapidly along the gradient of increasing C addition in plants with high Ellenberg N values than in plants with low Ellenberg N values. Soil C addition not only limits plant growth through a reduction in available nutrients, but also reduces mycorrhizal colonization of plant roots. The effect of C addition on mycorrhizal colonization varies among plant functional groups, with legumes experiencing an overproportional reduction in AM fungal colonization along the gradient of increasing C addition. We therefore propose that for a better understanding of vegetation succession on set-aside fields one may consider the interrelationship between plant growth, soil nutrient availability and mycorrhizal colonization of plant roots.     

Plant colonization windows in a mesic old field succession

Abstract. Closed canopy vegetation often prevents the colonization of plant species. Therefore the majority of plant species are expected to appear at the initial phase of post‐agricultural succession in mesic forest environment with moderate levels of resources. This hypothesis was tested with data from the Buell‐Small Successional Study, NJ, USA, one of the longest continuous fine‐scale studies of old‐field succession. The study started in 1958, including old fields with different agricultural histories, landscape contexts, and times of abandonment. In each year of the study, the cover values of plant species were recorded in 48 permanent plots of 1 m² in each field. We analysed the temporal patterns of colonization at plot scale and related these to precipitation data and other community characteristics. The number of colonizing species decreased significantly after ca. 5 yr, coinciding with the development of a continuous canopy of perennial species. However, species turnover remained high throughout the whole successional sequence. The most remarkable phenomenon is the high inter‐annual variation of all studied characteristics. We found considerable temporal collapses of vegetation cover that were synchronized among fields despite their different developmental stages and distinctive species compositions. Declines of total cover were correlated with drought events. These events were associated with peaks of local species extinctions and were followed by increased colonization rates. The transitions of major successional stages were often connected to these events. We suggest that plant colonization windows opened by extreme weather events during succession offer optimum periods for intervention in restoration practice.     

The colonization of ball-clay ponds by macroinvertebrates and macrophytes

SUMMARY. 1. The rate and nature of colonization of ball-clay ponds by aquatic macrophytes and macroinvertebrates were studied by comparing communities present in a scries of ten ponds of similar dimensions, but of different ages, ranging from 6 months to 15 years.
2. Multivariate analyses of the biotic data distinguished between'neutral'and'acid'ponds; further analyses concentrated on the former type.
3. Initial colonization of neutral ponds was rapid with a predictable sequence of species arrival, probably reflecting short dispersal distances. Variations in macroinvertebrate invasion times were related to dispersal strategy and ability. As the pond aged, successional changes, linked with macrophyte colonization, produced a shift in dominance from algivores and predators towards epiphyton grazers and detritivores. Some evidence of succession of species within invertebrate orders was found.
4. Low pH affected colonization principally by preventing the establishment of acid-intolerant immigrant species, but also by retarding succession.
5. It is concluded that non-interactive models of insular colonization, with constant immigration and extinction rates, are of limited applicability to pond colonization in temperate areas.     

The effects of enrichment on the dynamics of apparent competitive interactions in stage-structured systems

In the absence of other limiting factors, assemblages in which species share a common, effective natural enemy are not expected to persist. Although a variety of mechanisms have been postulated to explain the coexistence of species that share natural enemies, the role of productivity gradients has not been explored in detail. Here, we examine how enrichment can affect the outcome of apparent competition. We develop a structured resource/consumer/natural enemy model in which the prey are exposed to attacks during a vulnerable life phase, the length of which depends on resource availability. With a single prey species, the model exhibits the "paradox of enrichment," with unstable dynamics at high levels of resource productivity. We extend this model to consider two prey species linked by a shared predator, each with their own distinct resource base. We derive invasion and stability conditions and examine how enrichment influences prey species exclusion and coexistence. Contrary to expectations from simpler, prey-dependent models, apparent competition is not necessarily strong at high productivity, and prey species coexistence may thus be more likely in enriched environments. Further, the coexistence of apparent competitors may be facilitated by unstable dynamics. These results contrast with the standard theory that apparent competition in productive environments leads to nonpersistent interactions and that coexistence of multispecies interactions is more likely under equilibrial conditions.     

The role of preadaptation,propagule pressure and competition in the colonization of new habitats

To successfully colonize new habitats, organisms not only need to gain access to it, they also need to cope with the selective pressures imposed by the local biotic and abiotic conditions. The number of immigrants, the preadaptation to the local habitat and the presence of competitors are important factors determining the success of colonization. Here, using two experimental set-ups, we studied the effect of interspecific competition in combination with propagule pressure and preadaptation on the colonization success of new habitats. Our model system consisted of tomato plants (the novel habitat), the two-spotted spider mite Tetranychus urticae as our focal species and the red spider mite Tetranychus evansi as a competitor. Our results show that propagule pressure and preadaptation positively affect colonization success. More successful populations reach larger final population sizes either by having higher per capita growth rates (due to preadaptation effects) or by starting a population with a larger number of individuals. Although populations are more successful colonizing non-competitive environments than competitive ones, propagule pressure and preadaptation counteract the negative effects of competition, promoting colonization success. Our study shows the importance of propagule pressure and preadaptation for successful colonization of new habitats by providing the ability to cope with both the exigencies of new environments and the community context.     

Woody plant colonization in an experimentally fragmented landscape

The pattern of distribution and abundance of woody plants colonizing old fields is influenced by landscape spatial features, in particular, by the distance from the old field to propagule sources and the size of the habitat patches undergoing succession. Colonization is also influenced by species life history traits, such as dispersal mode, growth form, and fecundity. As part of a long-term project studying effects of habitat fragmentation on secondary succession at the prairie-forest ecotone, we have examined the colonization patterns of early-successional woody plants in an experimentally fragmented old field, with emphasis on the three woody species [Cornus drummondii C. A. Mey (rough-leaved dogwood), Ulmus rubra Muhl. (slippery elm), and Juniperus virginiana L. (red cedar)], which currently dominate the woody community on the site. The shapes of the colonization curve (proportion of colonized quadrats vs time) differed between C. drummondii and U. rubra. The rate of colonization by C. drummondii showed a pattern of acceleration after its initial colonization, consistent with rapid in situ recruitment from clonal growth and early seed production. By contrast, colonization by U. rubra fits a roughly linear pattern, consistent with recruitment only from external propagule sources. For both C. drummondii and U. rubra, density is currently greater in large patches than in small patches. No patch size difference was found for J. virginiana. The stern density of both C. drummondii and U. rubra exponentially decreased with distance to external propagule sources. The negative exponential pattern of U. rubra (wind-dispersed) with distance is sharper than that of C drummondii (bird-dispersed). Moreover, the amount of spatial variation in density explained by distance to source is greater on small patches. Our results highlight the importance of life history traits of colonizing species and spatial aspects of habitat during succession.     

Non-random processes determine the colonization of groundwater sediments by microbial communities in a pristine porous aquifer

Sediments accommodate the dominating share of groundwater microbiomes, however the processes that govern the assembly and succession of sediment-attached microbial communities in groundwater aquifers are not well understood. To elucidate these processes, we followed the microbial colonization of sterile sediments in in situ microcosms that were exposed to groundwater for almost 1 year at two distant but hydrologically connected sites of a pristine, shallow, porous aquifer. Our results revealed intriguing similarities between the community succession on the newly-colonized sediments and succession patterns previously observed for biofilms in other more dynamic aquatic environments, indicating that the assembly of microbial communities on surfaces may be governed by similar underlying mechanisms across a wide range of different habitats. Null model simulations on spatiotemporally resolved 16S rRNA amplicon sequencing data further indicated selection of specific OTUs rather than random colonization as the main driver of community assembly. A small fraction of persistent OTUs that had established on the sediments during the first 115 days dominated the final communities (68%–85%), suggesting a key role of these early-colonizing organisms, in particular specific genera within the Comamonadaceae and Oxalobacteraceae, for community assembly and succession during the colonization of the sediments. Overall, our study suggests that differences between planktonic and sediment-attached communities often reported for groundwater environments are not the result of purely stochastic events, but that sediment surfaces select for specific groups of microorganisms that assemble over time in a reproducible, non-random way.     

Benthic colonization and succession on temperate sublittoral rocky cliffs

Patterns of benthic colonization and succession were investigated on a temperate rocky cliff (Aegean Sea, Eastern Mediterranean). Cement and ceramic panels deployed on the rocky substratum at 25-30 m depth were sampled every 3 months over a 2-year period yielding 28 floral and 156 animal species. Diversity, cover and abundance had low values at short immersion periods and increased at the long ones. The vagile fauna responded to the increasing habitat complexity offered by the development of algal turfs and sessile species, and the community structure remained highly dynamic. Unlike the duration of immersion, the type of substratum didn't affect species colonization except for decapods that showed a preference for cement panels. Three to four stages of succession were recorded over immersion periods, according to vagile and sessile biota, respectively, during which species richness and abundance increased. The structure of the developed communities on both artificial materials differed from the natural algal-dominated benthic community of the same area, suggesting that the recovery of rocky shore communities on temperate cliffs is time consuming.     

Role of colonization in spatio-temporal patchiness of microgastropods in coralline turf habitat

The ability of benthic macrofauna to disperse and colonize new habitats throughout their life may contribute substantially to small-scale patchiness in abundances in different habitats. Microgastropods in coralline turf on rocky shores in Australia are very patchy in abundance at different spatio-temporal scales. They therefore represent an ideal assemblage for testing hypotheses about processes of colonization. Patterns and rates of colonization of 10 species of microgastropods were investigated in one intertidal habitat (coralline turf) in Botany Bay, New South Wales, Australia, using artificial substrata, which are considered to be a good mimic of natural coralline turfs. The experiment was designed to test the hypotheses that (1) patterns (abundance of colonizing individuals) and mode (juveniles vs. adults) of colonization depends on the proximity of a patch to a potential source of dispersing colonists (i.e. patch of natural coralline turf), (2) different species show different rates of colonization, and (3) patterns of succession are not repeatable among different patches of natural algae. Seven different plots (natural patches of coralline turf) were randomly chosen in the area of study. Artificial units (called patches) were placed at different distances around each plot (within the plot and 0, 50 or 100 cm away from the edge). Samples were collected 6, 13, and 27 days after the experiment started. Colonization was rapid (i.e. within 6 days) for most species. Pattern (number of individuals) and mode of colonization (adults vs. juveniles) varied among algal plots. Most species responded differently across patches causing no consistent patterns of colonization. Furthermore, patterns of colonization of artificial units were not always synchronous with, or in the same direction as, changes in abundances in the nearest algal plots, which themselves showed no consistent spatio-temporal pattern.     

Predation,apparent competition,and the structure of prey communities

It is argued that alternate prey species in the diet of a food-limited generalist predator should reduce each other's equilibrial abundances, whether or not they directly compete. Such indirect, interspecific interactions are labeled apparent competition. Two examples are discussed in which an observed pattern of habitat segregation was at first interpreted as evidence for direct competition, but later interpreted as apparent competition resulting from shared predation. In order to study the consequences of predator-mediated apparent competition in isolation from other complicating factors, a model community is analyzed in which there is no direct interspecific competition among the prey. An explicit necessary condition for prey species coexistence is derived for the case of one predator feeding on many prey species. This model community has several interesting properties: (1) Prey species with high relative values for a parameter $\text{r}\text{a}$ are “keystone” species in the community; (2) prey species can be excluded from the community by “diffuse” apparent competition; (3) large changes in the niche breadth of the predator need not correspond to large changes in predator density; (4) the prey trophic level as a whole is regulated by the predator, yet each of its constituent species is regulated by both the predator and available resources; (5) increased productivity may either increase, decrease, or leave unchanged the number of species in the community; (6) a decrease in density-independent mortality may decrease species diversity. These conclusions seem to be robust to changes in the prey growth equations and to the incorporation of predator satiation. By contrast, adding prey refugia or predator switching to the model weakens these conclusions. If the predator can be satiated or switched, the elements a_ij comprising the community matrix may have signs opposite the long-term effect of j upon i. The effect of natural selection upon prey species coexistence is discussed. Unless r_i, K_i, and a_i are tightly coupled, natural selection within prey species i will tend to decrease the equilibrial abundance of species j.     

Potential tree species extinction,colonization and recruitment in Afromontane forest relicts

Tree species regeneration determines future forest structure and composition, but is often severely hampered in small forest relicts. To study succession, long-term field observations or simulation models are used but data, knowledge or resources to run such models are often scarce in tropical areas. We propose and implement a species accounting equation, which includes the co-occurring events extinction, colonization and recruitment and which can be solved by using data from a single inventory. We solved this species accounting equation for the 12 remaining Afromontane cloud forest relicts in Taita Hills, Kenya by comparing the tree species present among the seedling, sapling and mature tree layer in 82 plots. A simultaneous ordination of the seedling, sapling and mature tree layer data revealed that potential species extinctions, colonizations and recruitments may induce future species shifts. On landscape level, the potential extinction debt amounted to 9% (7 species) of the regional species pool. On forest relict level, the smallest relicts harbored an important proportion of the tree species diversity in the regeneration layer. The average potential recruitment credit, defined as species only present as seedling or sapling, was 3 and 6 species for large and small forest relicts, while the average potential extinction debt was 12 and 4 species, respectively. In total, both large and small relicts are expected to lose approximately 20% of their current local tree species pool. The species accounting equations provide a time and resource effective tool and give an improved understanding of the conservation status and possible future succession dynamics of forest relicts, which can be particularly useful in a context of participatory monitoring.     

Litter and soil arthropod colonization in reforested semi‐deciduous seasonal Atlantic forests

The recovery of soil ecological processes during the restoration of tropical forests is greatly influenced by arthropods that live in the litter and soil. However, these communities present complex dynamics, and their colonization patterns are not well understood. In this study, we examined the response patterns of litter and soil arthropods to the ecosystem regeneration process by assessing reforestation sites from two regions of São Paulo State, Brazil, and we compared the data obtained from these sites with data from mature forests. We assessed the arthropod communities using similarity indices and high‐level taxa abundance, with the level of forest succession and the locations of the restoration areas as factors. Forest succession correlated with the species composition as communities from the reforestation sites gradually became more similar to communities from the mature forests, while their quantitative patterns were minimally related. Forest maturation positively affected the richness of the litter community and the abundance of some minor groups, such as Protura, Diplura, and Symphyla. The region influenced the species composition but did not influence the manner in which the communities changed during the maturation process. We also found a convergent soil colonization pattern as arthropod communities from different sites became more similar during forest succession. This finding is consistent with both empirical data and theoretical predictions from the specialized literature, although the subject has been poorly explored until now. We conclude that reforestation allows the colonization of soil and litter fauna in a biased manner.     

Exploring tree species colonization potentials using a spatially explicit simulation model: implications for four oaks under climate change

Climate change impacts tree species differentially by exerting unique pressures and altering their suitable habitats. We previously predicted these changes in suitable habitat for current and future climates using a species habitat model (DISTRIB) in the eastern United States. Based on the accuracy of the model, the species assemblages should eventually reflect the new quasi‐equilibrium suitable habitats (~2100) after accounting for the lag in colonization. However, it is an open question if and when these newly suitable habitats will be colonized under current fragmented landscapes and realistic migration rates. To evaluate this, we used a spatially explicit cell‐based model (SHIFT) that estimates colonization potentials under current fragmented habitats and several estimates of historical migration rates at a 1 km resolution. Computation time, which was previously the biggest constraint, was overcome by a novel application of convolution and Fast Fourier Transforms. SHIFT outputs, when intersected with future suitable habitats predicted by DISTRIB, allow assessment of colonization potential under future climates. In this article, we show how our approach can be used to screen multiple tree species for their colonization potentials under climate change. In particular, we use the DISTRIB and SHIFT models in combination to assess if the future dominant forest types in the north will really be dominated by oaks, as modelled via DISTRIB. Even under optimistic scenarios, we conclude that only a small fraction of the suitable habitats of oaks predicted by DISTRIB is likely to be occupied within 100 years, and this will be concentrated in the first 10–20 km from the current boundary. We also show how DISTRIB and SHIFT can be used to evaluate the potential for assisted migration of vulnerable tree species, and discuss the dynamics of colonization at range limits.     

The role of innate immune responses in the outcome of interspecies competition for colonization of mucosal surfaces

Since mucosal surfaces may be simultaneously colonized by multiple species, the success of an organism may be determined by its ability to compete with co-inhabitants of its niche. To explore the contribution of host factors to polymicrobial competition, a murine model was used to study the initiation of colonization by Haemophilus influenzae and Streptococcus pneumoniae. Both bacterial species, which occupy a similar microenvironment within the nasopharynx, persisted during colonization when given individually. Co-colonization, however, resulted in rapid clearance of S. pneumoniae from the upper respiratory tract, associated with increased recruitment of neutrophils into paranasal spaces. Systemic depletion of either neutrophil-like cells or complement was sufficient to eliminate this competitive effect, indicating that clearance was likely due to enhanced opsonophagocytic killing. The hypothesis that modulation of opsonophagocytic activity was responsible for host-mediated competition was tested using in vitro killing assays with elicited neutrophil-like cells. Components of H. influenzae (but not S. pneumoniae) stimulated complement-dependent phagocytic killing of S. pneumoniae. Thus, the recruitment and activation of neutrophils through selective microbial pattern recognition may underlie the H. influenzae-induced clearance of S. pneumoniae. This study demonstrates how innate immune responses may mediate competitive interactions between species and dictate the composition of the colonizing flora.     

Mathematical models of cell colonization of uniformly growing domains

During the development of vertebrate embryos, cell migrations occur on an underlying tissue domain in response to some factor, such as nutrient. Over the time scale of days in which this cell migration occurs, the underlying tissue is itself growing. Consequently cell migration and colonization is strongly affected by the tissue domain growth. Numerical solutions for a mathematical model of chemotactic migration with no domain growth can lead to travelling waves of cells with constant velocity; the addition of domain growth can lead to travelling waves with nonconstant velocity. These observations suggest a mathematical approximation to the full system equations, allowing the method of characteristics to be applied to a simplified chemotactic migration model. The evolution of the leading front of the migrating cell wave is analysed. Linear, exponential and logistic uniform domain growths are considered. Successful colonization of a growing domain depends on the competition between cell migration velocity and the velocity and form of the domain growth, as well as the initial penetration distance of the cells. In some instances the cells will never successfully colonize the growing domain. These models provide an insight into cell migration during embryonic growth, and its dependence upon the form and timing of the domain growth.     

Convergence of community composition during secondary succession on Zokor rodent mounds on the Tibetan Plateau

Aims The community succession theory is much debated in ecology. We studied succession on Zokor rodent mounds on the Tibetan Plateau to address several fundamental questions, among them: (i) During secondary succession, does the community composition converge towards one community state or multiple states depending on the initial colonization? (ii) Do mound communities located in different background communities exhibit different assembly trajectories?Methods In a sub-alpine meadow, we investigated a total of 80 mound communities at several successional stages in three different background communities resulting from different management histories and compared their changes in species composition. The distribution of plant communities over time was analyzed with quantitative classification and ordination methods. The co-occurrence patterns of species were evaluated at each successional stage, and the degree of convergence/divergence among communities was obtained by calculating two beta-diversity indices.Important findings During secondary succession, species richness of mound communities changed over time, and this change was dependent on the background community. Five life-form groups exhibited different dynamic patterns in species richness and plant cover. Community composition and the degree of species co-occurrence between communities increased over time since disturbance. There was much variation in species composition at earlier stages of succession, but communities on older mounds became more similar to each other and to their surrounding vegetation over the course of secondary succession. Post-disturbance succession of Zokor mound communities transitioned from 'multiple alternative states' to 'background-based deterministic community assembly' over time. Tradeoffs between competition and colonization, as well as the characteristics of different life-forms and mass effects within a limited species pool are the mechanisms responsible for convergence of mound communities.     

Evaluation of assisted colonization strategies under global change for a rare,fire‐dependent plant

As a clear consensus is emerging that habitat for many species will dramatically reduce or shift with climate change, attention is turning to adaptation strategies to address these impacts. Assisted colonization is one such strategy that has been predominantly discussed in terms of the costs of introducing potential competitors into new communities and the benefits of reducing extinction risk. However, the success or failure of assisted colonization will depend on a range of population‐level factors that have not yet been quantitatively evaluated – the quality of the recipient habitat, the number and life stages of translocated individuals, the establishment of translocated individuals in their new habitat and whether the recipient habitat is subject to ongoing threats all will play an important role in population persistence. In this article, we do not take one side or the other in the debate over whether assisted colonization is worthwhile. Rather, we focus on the likelihood that assisted colonization will promote population persistence in the face of climate‐induced distribution changes and altered fire regimes for a rare endemic species. We link a population model with species distribution models to investigate expected changes in populations with climate change, the impact of altered fire regimes on population persistence and how much assisted colonization is necessary to minimize risk of decline in populations of Tecate cypress, a rare endemic tree in the California Floristic Province, a biodiversity hotspot. We show that assisted colonization may be a risk‐minimizing adaptation strategy when there are large source populations that are declining dramatically due to habitat contractions, multiple nearby sites predicted to contain suitable habitat, minimal natural dispersal, high rates of establishment of translocated populations and the absence of nonclimatic threats such as altered disturbance regimes. However, when serious ongoing threats exist, assisted colonization is ineffective.     

Extinction, colonization, and species occupancy in tidepool fishes

Despite the increasing sophistication of ecological models with respect to the size and spatial arrangement of habitat, there is relatively little empirical documentation of how species dynamics change as a function of habitat size and the fraction of habitat occupied. In an assemblage of tidepool fishes, I used maximum-likelihood estimation to test whether models which included habitat size provided a better fit to empirical data on extinction and colonization probabilities than models that assumed constant probabilities over all habitats. I found species differences in how extinction and colonization probabilities scaled with habitat size (and hence local population size). However, there was little evidence for a relationship between extinction and colonization probabilities and the fraction of occupied tidepools, as assumed in simple metapopulation models. Instead, colonization and extinction were independent of the fraction of occupied tidepools, favoring a MacArthur-Wilson island-mainland model. When I incorporated declines in extinction probability with tidepool volume in a simple simulation model, I found that predicted occupancy could change greatly, especially when colonization was low. However, the predicted fraction of occupied patches in the simulation model changed little when I incorporated the range of values reported here for extinction and colonization and the rate at which they scale with habitat size. Quantifying extinction and colonization patterns of natural populations is fundamental to understanding how species are distributed spatially and whether metapopulation models of species occupancy provide explanatory power for field populations. Received: 14 March 1997 / Accepted: 21 September 1997