Metapopulation dynamics of the bog fritillary butterfly: comparison of demographic parameters and dispersal between a continuous and a highly fragmented landscape

We investigated the effects of habitat loss and fragmentation on population functioning. We compared demography (daily and total population sizes) and dispersal (dispersal rate and dispersal kernels) of the bog fritillary butterfly in two 6-km² landscapes differing in their degree of fragmentation. In 2000, we conducted a Capture-Mark-Recapture experiment in a highly fragmented system in the marginal part of the species distribution (Belgium) and in a more continuous system in the central part of its distribution (Finland). A total of 293 and 947 butterflies were marked with 286 and 190 recapture events recorded in the fragmented and the continuous system respectively. Our results suggest that habitat loss and fragmentation affect dispersal more than demography. Although density was lower in the continuous system, it remains in the yearly range of variation observed on 10 generations in the fragmented system. However, in the fragmented system, the dispersal rate dropped drastically (39 vs. 64%) and females moved longer distances. Patch area had a significant effect on migration in the fragmented system only. From our results, we propose the definition of a new parameter, the minimal patch area (MPA) needed to establish a local population in highly fragmented landscapes.This revised version was published online in May 2005 with corrections to the Cover Date.     

A controlled,hierarchical study of habitat fragmentation: responses at the individual,patch, and landscape scale

We compared the performance of individuals and whole populations of meadow voles, Microtus pennsylvanicus, within and between experimentally created habitat fragments of three sizes (1.0, 0.25, and 0.0625 ha) and between a 20-ha fragmented and a 20-ha continuous habitat landscape. We recorded 10,020 captures of 3946 individuals over 17 censuses between June 1993 and October 1994. Five demographic parameters showed significantly different population responses between the two landscapes but no difference in tests comparing fragment size: i.e., mean and peak population densities (the latter, in each of the two growing seasons) averaged 149 to 172% higher, population growth rate averaged 219% higher, and adult recruitment 170% higher in fragmented than in the continuous control landscape. Observations at the individual level (body sizes, rates of reproduction, residence times) suggested that these landscape differences involved enhanced performance of adult females associated with edge habitats rather than differential immigration or emigration. If this turns out to be a common response to fragmentation, the detection of such responses will be greater when comparing fragmented and unfragmented landscapes with qualitatively different structure than for fragments of varied size with differing proportions of edge. That responses to habitat fragmentation may be more evident at the very small (individual) and very large (landscape) scales, but may be obscured at the intermediate spatial scale of fragments, is a proposition that clearly requires more attention.     

Migration rates of grassland plants along corridors in fragmented landscapes assessed with a cellular automation model

This study investigated the efficacy of linear landscape elements in fragmented landscapes as corridors for perennial grassland species with short-range seed dispersal. Corridors are assumed to be essential for the persistence of metapopulations in fragmented landscapes, but it is unclear to what extent linear landscape elements such as ditch banks and road verges can function as corridors for those species. The principal factors that determine the rate of migration through corridors include the width and habitat quality of patches within a corridor (expressed as the population growth rate λ) and the dispersal capacity of plants (expressed as the slope α of the relationship between seed number and log-distance). A cellular automation model was used to simulate the effects of the principal factors on the rate of migration. Simulations with different levels of the principal factors showed highly significant and positive main effects of dispersal capacity, habitat quality and width of corridors on the migration rate. Significant interactions existed between dispersal capacity × width and dispersal capacity × habitat quality (p<0.0001), indicating that the effects of width and habitat quality depended on the dispersal capacity. In narrow corridors most of the dispersed seeds were deposited outside the corridor, which significantly reduced migration rates, especially for species with long-range dispersal (α=−0.4). In wide corridors (up to 20 m), seed losses were much smaller and migration rates approximated those of continuous habitats. The contribution of the few long-range seeds to the rate of migration was significant when habitat quality was high (population growth rates up to 2.5). However, in all simulations migration rates were very low,i.e.<5 m/yr. It is concluded that linear landscape elements are not effective corridors in fragmented landscapes for plants with short-range seed dispersal, because migration rates are low (<5 m/yr), landscape elements vary in the percentage of high quality patches, and refugia and suitable habitat patches are frequently several kilometres apart, making a cohesive infrastructure of corridors for plants elusive. It is argued that the best way to conserve endangered plant species that encounter dispersal barriers is to harvest seeds from nearby source populations and introduce them as suitable habitats.     

Metapopulation theory and habitat fragmentation: a review of holarctic breeding bird studies

Metapopulations are conceived as spatially structured populations consisting of distinct units (subpopulations), separated by space or barriers, and connected by dispersal movements. Metapopulations characteristically demonstrate a turnover of local populations going extinct and becoming re-established, resulting in a distribution pattern that shifts over time. Metapopulation theory is used to analyse the effects of habitat fragmentation on birds in the temperate zone, integrating various explanations for the paucity of species in isolated ecotopes.There is some evidence that turnover of local populations occurs in fragmented systems. A few studies based on time series demonstrate the local extinction rate to be related to the size of the habitat fragment, whereas the recolonization rate depends on the degree of isolation. Most evidence comes from short-term pattern studies in which the probability of occurrence was found to depend on the size of habitat fragments, on their relative position in the landscape and on the density of corridors lowering the landscape resistance. These data are consistent with predictions from metapopulation theory. However, almost all investigations consider wood fragmentation in agricultural landscapes, and there is a great need for studies in naturally fragmented landscapes as well as for studies focussing on other, less predictable, habitat types.     

Analyzing the effect of stepping stones on target patch colonisation in structured landscapes for Eurasian lynx

With habitat loss and fragmentation having become two of the major threats to the viability of species, the question of how to manage landscapes for species conservation has attracted much attention. In this context, the planning of stepping stones has been proposed to increase connectivity in fragmented landscapes. We present a simulation study with a neutral landscape approach to assess the effects of stepping stones on colonization success. To that end, we used a spatially explicit, calibrated population model of the European lynx (Lynx lynx) coupled with structured landscapes, in which we could control the landscape parameters of dispersal habitat coverage and contagion, as well as the number and size of stepping stones available for breeding. In general, we found that colonization success increased with increasing habitat coverage but decreased with increasing habitat contagion, while the introduction of stepping stones had significant effects in critical situations. Especially at low to medium dispersal habitat coverage and high disperser mortality, stepping stones had a positive effect on colonization success when they were large enough to produce new dispersers, but negative effects when they were small and located in a way that dispersers would be distracted from more suitable breeding habitat patches. The latter clearly constituted a shading effect and argues for a thorough consideration of the trade-offs related to stepping stone size and location when implementing stepping stones as a conservation measure, especially when the number of individuals of conservation concern is low.     

Effects of habitat fragmentation on meadow vole (Microtus pennsylvanicus) population dynamics in experiment landscape patches

This study examined the effects of habitat fragmentation on meadow vole (Microtus pennsylvanicus) population dynamics in experimental landscape patches. The study was conducted from May–November 1993 at the Miami University Ecology Research Center. Eight 0.1-ha small mammal enclosures were used. Four enclosures contained a 160 m² nonfragmented patch and four enclosures contained four 40 m² fragmented patches. Thus, each treatment was replicated 4 times in a systematic research design. The patches in both treatments contained high-quality habitat surrounded by low-quality matrix. Six pairs of adult meadow voles were released in each enclosure on 27 May 1993. Populations were monitored by live-trapping and radio-telemetry methods. Significantly greater densities of female voles were found during October in the fragmented treatment compared to the nonfragmented treatment. Also, significantly more females than males were found in the fragmented treatment compared to the nonfragmented treatment for the total study period. Significantly more subadult and juvenile males were found in the matrix versus the patch of the nonfragmented treatment compared to the fragmented treatment. Males in the fragmented treatment had significantly greater mean home range size than males or females in the nonfragmented treatment. There appears to exist a relationship between patch fragmentation and the social structure of meadow vole populations; this relationship appears to function as a population regulatory mechanism.     

Matrix quality and habitat configuration interactively determine functional connectivity in a widespread bush cricket at a small spatial scale

Unlike rare or specialised species, widespread abundant species have often been neglected when studying effects of habitat fragmentation. However, recently, it was shown that in the widespread abundant bush cricket Pholidoptera griseoaptera gene flow becomes restricted when the share of suitable habitat dropped below a threshold of 20% at the landscape scale. Here, using the same highly fragmented landscape, we studied the impact of habitat configuration and matrix quality on genetic variation and population differentiation of P. griseoaptera at a small spatial scale. We investigated four clusters of three populations that were either disconnected or connected and had either low quality (arable land) or high quality (grassland) matrix. The number of alleles was significantly lower in disconnected than in connected clusters, irrespective of matrix quality. Genetic differentiation was equally high in the two disconnected clusters and in the connected cluster with low quality matrix (G _ST ≥ 0.030; D ≥ 0.082), whereas it was significantly reduced when connected habitats were embedded in a high quality grassland matrix (G _ST = 0.004; D = 0.011). Analyses of least-cost paths showed that grassy landscape elements in fact represent high quality matrix, but that linear grassy margins are costly for dispersal. The effect of habitat configuration on genetic diversity may be explained by lower effective population sizes in disconnected habitats. The fact that only the connected populations in high quality matrix were not differentiated indicates that landscape management should simultaneously consider habitat configuration and matrix quality to effectively promote small and dispersal-limited species, also at small spatial scales.     

Interactions between plant life span, seed dispersal capacity and fecundity determine metapopulation viability in a dynamic landscape

Classical metapopulation models do not account for temporal changes in the suitability of habitat patches. In reality, however, the carrying capacity of most habitat types is not constant in time due to natural succession processes. In this study, we modeled plant metapopulation persistence in a successional landscape with disappearing and emerging habitat patches, based on a realistic dune slack landscape at the Belgian–French coast. We focused on the effects of the variation of different plant traits on metapopulation persistence in this changing landscape. Therefore, we used a stage based stochastic metapopulation model implemented in RAMAS/Metapop, simulating a large variation in plant traits but keeping landscape characteristics such as patch turnover rate and patch lifespan constant. The results confirm the conclusions of earlier modeling work that seed dispersal distance and seed emigration rate both have an important effect on metapopulation persistence. We also found that high population growth rate or high recruitment considerably decreased the extinction risk of the metapopulation. Additionally, a long plant life span had a strong positive effect on metapopulation persistence, irrespective of the plant's dispersal capacity and population growth rate. Plant species that invest in life span require less investment in offspring and dispersal capacity to avoid extinction, even in dynamic landscapes with deterministic changes in habitat quality. Moreover, metapopulations of long-lived plant species were found to be much less sensitive to high levels of environmental stochasticity than short-lived species.     

Dispersal success on fractal landscapes: a consequence of lacunarity thresholds

Habitat fragmentation is expected to disrupt dispersal, and thus we explored how patch metrics of landscape structure, such as percolation thresholds used to define landscape connectivity, corresponded with dispersal success on neutral landscapes. We simulated dispersal as either a purely random process (random direction and random step lengths) or as an area-limited random walk (random direction, but movement limited to an adjacent cell at each dispersal step) and quantified dispersal success for 1000 individuals on random and fractal landscape maps across a range of habitat abundance and fragmentation. Dispersal success increased with the number of cells a disperser could search (m), but poor dispersers (m<5) searching via area-limited dispersal on fractal landscapes were more successful at locating suitable habitat than random dispersers on either random or fractal landscapes. Dispersal success was enhanced on fractal landscapes relative to random ones because of the greater spatial contagion of habitat. Dispersal success decreased proportionate to habitat loss for poor dispersers (m=1) on random landscapes, but exhibited an abrupt threshold at low levels of habitat abundance (p<0.1) for area-limited dispersers (m<10) on fractal landscapes. Conventional metrics of patch structure, including percolation, did not exhibit threshold behavior in the region of the dispersal threshold. A lacunarity analysis of the gap structure of landscape patterns, however, revealed a strong threshold in the variability of gap sizes at low levels of habitat abundance (p<0.1) in fractal landscapes, the same region in which abrupt declines in dispersal success were observed. The interpatch distances or gaps across which dispersers must move in search of suitable habitat should influence dispersal success, and our results suggest that there is a critical gap-size structure to fractal landscapes that interferes with the ability of dispersers to locate suitable habitat when habitat is rare. We suggest that the gap structure of landscapes is a more important determinant of dispersal than patch structure, although both are ultimately required to predict the ecological consequences of habitat fragmentation.     

Population genetic structure and landscape connectivity of the Eastern Yellowbelly Racer (<Emphasis Type="Italic">Coluber constrictor flaviventris</Emphasis>) in the contiguous tallgrass prairie of northeastern Kansas,USA

The tallgrass prairie of North America has undergone widespread habitat loss and fragmentation (<4% remains). The Flint Hills region of Kansas and Oklahoma is the largest tallgrass prairie remaining and therefore provides an opportunity to study the population genetic structure of grassland species in a relatively contiguous landscape and set a baseline for evaluating changes when the habitat is fragmented. We adopted a landscape genetics approach to identify how landscape structure affected dispersal, population genetic structure, and landscape connectivity of the Eastern Yellowbelly Racer (Coluber constrictor flaviventris) across a 13,500-km² landscape in northeastern Kansas, USA. The racer population had high allelic diversity, high heterozygosity, and was maintaining migration-drift equilibrium. Autocorrelation between genetic and geographic distance revealed that racers exhibited restricted dispersal within 3 km, and isolation-by-distance. Significant isolation-by-distance occurred at broad regional scales (>100 km), but because of sufficient gene flow between locations, we were unable to define discrete subpopulations using Bayesian clustering analyses. Resistance distance, which considers the permeability of habitats, did not explain significant variation in genetic distance beyond Euclidean distance alone, suggesting that racers are not currently influenced by landscape composition. In northeastern Kansas, racers appear to be an abundant and continuously distributed snake that perceives the landscape as well connected with no cover type currently impeding snake dispersal or gene flow.     

Patterns of habitat occupancy, genetic variation and predicted movement of a flightless bush cricket, Pholidoptera griseoaptera, in an agricultural mosaic landscape

Habitat fragmentation has been generally regarded detrimental to the persistence of many species, especially those with limited dispersal abilities. Yet, when exactly habitat elements become functionally disconnected very much depends on the dispersal ability of a species in combination with the landscape’s composition in which it occurs. Surprisingly, for many small and ground-walking generalists knowledge at what spatial scale and to what extent landscape structure affects dispersal is very scarce. Because it is flightless, the bush cricket Pholidoptera griseoaptera may be regarded susceptible to fragmentation. We applied habitat occupancy surveys, population genetic analyses and movement modelling to investigate the performance of P. griseoaptera in an agricultural mosaic landscape with suitable habitat patches of varying size and isolation. Despite its presumed dispersal limitation we could show that P. griseoaptera occupied the majority of suitable habitats, including small and isolated patches, showed a very low and non-significant genetic differentiation (F _ST = 0.0072) and, in the model, managed to colonize around 73% of all suitable habitat patches within one generation under weak and strong landscape-effect scenarios. We conclude that P. griseoaptera possesses the behavioural attributes (frequent inter-patch dispersal) necessary to persist in this landscape characterized by a patchy distribution of habitat elements. Yet, sound recommendations to landscape planning and conservation require more research to determine whether this represents a general behaviour of the species or a behavioural adaptation to this particular landscape.     

Independent effects of connectivity predict homing success by northern flying squirrel in a forest mosaic

Landscape composition and configuration, often termed as habitat loss and fragmentation, are predicted to reduce species population viability, partly due to the restriction of movement in the landscape. Unfortunately, measuring the effects of habitat loss and fragmentation on functional connectivity is challenging because these variables are confounded, and often the motivation for movement by target species is unknown. Our objective was to determine the independent effects of landscape connectivity from the perspective of a mature forest specialist—the northern flying squirrel (Glaucomys sabrinus). To standardize movement motivation, we translocated 119 squirrels, at varying distances (0.18–3.8 km) from their home range across landscapes representing gradients in both habitat loss and fragmentation. We measured the physical connectedness of mature forest using an index of connectivity (landscape coincidence probability). Patches were considered connected if they were within the mean gliding distance of a flying squirrel. Homing success increased in landscapes with a higher connectivity index. However, homing time was not strongly predicted by habitat amount, connectivity index, or mean nearest neighbour and was best explained as a simple function of sex and distance translocated. Our study shows support for the independent effects of landscape configuration on animal movement at a spatial scale that encompasses several home ranges. We conclude that connectivity of mature forest should be considered for the conservation of some mature forest specialists, even in forest mosaics where the distinction between habitat and movement corridors are less distinct.     

Plant strategies and agricultural landscapes: survival in spatially and temporally fragmented habitat

In agricultural landscapes many plant species are limited to the network of landscape elements that are not used for agricultural production. This habitat is fragmented in space and time due to anthropogenic, biotic and abiotic factors. Therefore, plant populations are spatially sub-divided and their persistence might be dependent on the spatial dynamics in the network of local populations. Dispersal characteristics and seed bank persistence are main determinants of colonization ability which in turn is a key determinant of metapopulation viability. We propose a conceptual model that relates plant population dynamics to habitat quality, configuration and dynamics. In this model, the habitat is arranged as a network of suitable and unsuitable patches,and the distribution of the patches is assumed to be dynamic in time. Based on dispersal and seed bank characteristics four plant strategies are distinguished:species having either long (> 100 m) or short (< 100m) distance dispersal and either a long (> 5 yr)or short (< 5 yr) term persistent seed bank. We expect that species with contrasting strategies have different survival probabilities in landscapes with contrasting habitat arrangement in space and time. We found few empirical studies for testing the hypotheses based on the model. Therefore the relation between landscapes and plant survival needs to be further explored,especially the quantitative aspects. We propose an iterative process of empirical and modelling research to determine this relation and to define management options for multifunctional farms in which biodiversity is one of the land use aims. This revised version was published online in July 2006 with corrections to the Cover Date.     

Landscape ecology as a theoretical basis for nature conservation

Conservation of representative biotopes, single species populations or biodiversity usually embraces two or more biotopes, and is often affected by surrounding croplands. The conclusions from landscape ecological studies can, therefore, offer important contributions to conservation, especially at early levels of landscape change or habitat fragmentation. Indicator and keystone species are useful for monitoring and managing fragmented biotopes, respectively. Communities as well as single species are affected by the juxtaposition of successional and climax biotopes, which influence climatic equability, seasonality, productivity and dispersal. Low levels of fragmentation may result in ill-functioning communities, and greater fragmentation may result in species losses and ultimately in the loss of whole communities. Fragmented habitats retain species with high reproductive and dispersal rates and generalized habitat selection. New combinations of interacting species will lead to trivialization of earlier habitat-specific interactions. Validation of these concepts was made with data from a Swedish research program on fragmented biotopes in production landscapes. General reserve selection and methods of management for preserving climax communities, single specialized species and high biodiversity are suggested.     

How do local habitat management and landscape structure at different spatial scales affect fritillary butterfly distribution on fragmented wetlands?

Habitat fragmentation, patch quality and landscape structure are important predictors for species richness. However, conservation strategies targeting single species mainly focus on habitat patches and neglect possible effects of the surrounding landscape. This project assesses the impact of management, habitat fragmentation and landscape structure at different spatial scales on the distribution of three endangered butterfly species, Boloria selene, Boloria titania and Brenthis ino. We selected 36 study sites in the Swiss Alps differing in (1) the proportion of suitable habitat (i.e., wetlands); (2) the proportion of potential dispersal barriers (forest) in the surrounding landscape; (3) altitude; (4) habitat area and (5) management (mowing versus grazing). Three surveys per study site were conducted during the adult flight period to estimate occurrence and density of each species. For the best disperser B. selene the probability of occurrence was positively related to increasing proportion of wetland on a large spatial scale (radius: 4,000 m), for the medium disperser B. ino on an intermediate spatial scale (2,000 m) and for the poorest disperser B. titania on a small spatial scale (1,000 m). Nearby forest did not negatively affect butterfly species distribution but instead enhanced the probability of occurrence and the population density of B. titania. The fen-specialist B. selene had a higher probability of occurrence and higher population densities on grazed compared to mown fens. The altitude of the habitat patches affected the occurrence of the three species and increasing habitat area enhanced the probability of occurrence of B. selene and B. ino. We conclude that, the surrounding landscape is of relevance for species distribution, but management and habitat fragmentation are often more important. We suggest that butterfly conservation should not focus only on a patch scale, but also on a landscape scale, taking into account species-specific dispersal abilities.     

Modelling the Effects of Dispersal and Landscape Configuration on Population Distribution and Viability in Fragmented Habitat

Landscape configuration and dispersal characteristics are major determinants of population distribution and persistence in fragmented habitat. An individual-based spatially explicit population model was developed to investigate these factors using the distribution of nuthatches in an area of eastern England as an example. The effects of immigration and increasing the area of breeding quality habitat were explored. Predictions were compared with observed population sizes in the study area. Our model combined a nuthatch population simulator based on individual behaviour with a grid-based representation of the landscape; nuthatch life cycle and immigration parameters were user selectable. A novel aspect of the model is user-selection of habitat perceptual range. Using a realistic set of parameters, the number of breeding pairs predicted by the model matched observed numbers. According to model simulations, the main cause of nuthatch scarcity in the study area was the inability of patches to support viable populations without immigration from elsewhere. Modelled habitat management, which increased breeding quality habitat in existing woods, lowered the threshold above which the study area population became self-sustaining. The existence of a large core habitat area was critical in producing a self-sustaining population in this landscape, the same area in dispersed small woods failed to sustain populations.     

The importance of integrating landscape ecology in habitat models: isolation-driven occurrence of north island robins in a fragmented landscape

Although the role of habitat fragmentation in species declines is well recognised, the effect of habitat quality on species distributions is often studied using presence–absence models that ignore metapopulation dynamics. We compared three approaches to model the presence–absence of North Island robins in 400 sites among 74 fragments of native forest in a 15,000-ha agricultural landscape in New Zealand. The first approach only considered local habitat characteristics, the second approach only considered metapopulation factors (patch size and isolation), and the third approach combined these two types of factors. The distribution of North Island robins was best predicted by patch isolation, as their probability of occurrence was negatively correlated with isolation from neighbouring patches and from the closest major forests, which probably acted as a source of immigrants. The inclusion of habitat factors gave only a slight increase in predictive power and indicated that robins were more likely to occur in areas with tall canopy, tall understory and low density of young trees. We modelled the effect of isolation using an index of functional patch connectivity based on dispersal behaviour of radio-tracked juveniles, and this functional index greatly improved the models in comparison to classical indices relying on Euclidean distances. This study highlights the need to incorporate functional indices of isolation in presence–absence models in fragmented landscapes, as species occurrence can otherwise be a misleading predictor of habitat quality and lead to wrong interpretations and management recommendations.     

Using an individual-based model to examine the roles of habitat fragmentation and behavior on predator–prey relationships in seagrass landscapes

Seagrasses, which form critical subtidal habitats for marine organisms worldwide, are fragmented via natural processes but are increasingly being fragmented and degraded by boating, fishing, and coastal development. We constructed an individual-based model to test how habitat fragmentation and loss influenced predator–prey interactions and cohort size for a group of settling juvenile blue crabs (Callinectes sapidus Rathbun) in seagrass landscapes. Using results from field studies suggesting that strong top-down processes influence the relationship between cannibalistic blue crab populations and seagrass landscape structure, we constructed a model in which prey (juvenile blue crabs) are eaten by mesopredators (larger blue crabs) which in turn are eaten by top-level predators (e.g., large fishes). In our model, we varied the following parameters within four increasingly fragmented seagrass landscapes to test for their relative effects on cohort size: juvenile blue crab (prey) predator avoidance response, hunting ability of mesopredators and predators, the presence of a top-level predator, and prey settlement routines. Generally, prey cohort size was maximized in the presence of top-level predators and when mesopredators and predators exhibited random searching behavior vs. directed hunting. Cohort size for stationary (tethered) prey was maximized in fragmented landscapes, which corresponds to results from field experiments, whereas mobile prey able to detect and avoid predators had higher survival in continuous landscapes. Prey settlement patterns had relatively small influences on cohort size. We conclude that the effects of seagrass fragmentation and loss on organisms such as blue crabs will depend heavily on behaviors of prey and predatory organisms and how these behaviors change with landscape structure.

     

Population dynamics and habitat connectivity affecting the spatial spread of populations – a simulation study

In this paper we show how the spatialconfiguration of habitat quality affects the spatial spread of apopulation in a heterogeneous environment. Our main result is thatfor species with limited dispersal ability and a landscape withisolated habitats, stepping stone patches of habitat greatlyincrease the ability of species to disperse. Our results showthat increasing reproductive rate first enables and thenaccelerates spatial spread, whereas increasing the connectivity has aremarkable effect only in case of low reproductive rates. Theimportance of landscape structure varied according to thedemographic characteristics of the population. To show this wepresent a spatially explicit habitat model taking into accountpopulation dynamics and habitat connectivity. The population dynamicsare based on a matrix projection model and are calculated on eachcell of a regular lattice. The parameters of the Leslie matrix dependon habitat suitability as well as density. Dispersal between adjacentcells takes place either unrestricted or with higher probability inthe direction of a higher habitat quality (restricted dispersal).Connectivity is maintained by corridors and stepping stones ofoptimal habitat quality in our fragmented model landscape containinga mosaic of different habitat suitabilities. The cellular automatonmodel serves as a basis for investigating different combinations ofparameter values and spatial arrangements of cells with high and lowquality.This revised version was published online in May 2005 with corrections to the Cover Date.