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.     

Habitat quality of source patches and connectivity in fragmented landscapes

Because spatial connectivity is critical to dispersal success and persistence of species in highly fragmented landscapes, the way that we envision and measure connectivity is consequential for biodiversity conservation. Connectivity metrics used for predictive modeling of spatial turnover and patch occupancy for metapopulations, such as with Incidence Function Models (IFM), incorporate distances to and sizes of possible source populations. Here, our focus is on whether habitat quality of source patches also is considered in these connectivity metrics. We propose that effective areas (weighted by habitat quality) of source patches should be better surrogates for population size and dispersal potential compared to unadjusted patch areas. Our review of a representative sample of the literature revealed that only 12.5% of studies incorporated habitat quality of source patches into IFM-type connectivity metrics. Quality of source patches generally was not taken into account in studies even if habitat quality of focal patches was included in analyses. We provide an empirical example for a metapopulation of a rare wetland species, the round-tailed muskrat (Neofiber alleni), demonstrating that a connectivity metric based on effective areas of source patches better predicts patch colonization and occupancy than a metric that used simple patch areas. The ongoing integration of landscape ecology and metapopulation dynamics could be hastened by incorporating habitat quality of source patches into spatial connectivity metrics applied to species conservation in fragmented landscapes.     

Metapopulation dynamics in a changing climate: Increasing spatial synchrony in weather conditions drives metapopulation synchrony of a butterfly inhabiting a fragmented landscape

Habitat fragmentation and climate change are both prominent manifestations of global change, but there is little knowledge on the specific mechanisms of how climate change may modify the effects of habitat fragmentation, for example, by altering dynamics of spatially structured populations. The long‐term viability of metapopulations is dependent on independent dynamics of local populations, because it mitigates fluctuations in the size of the metapopulation as a whole. Metapopulation viability will be compromised if climate change increases spatial synchrony in weather conditions associated with population growth rates. We studied a recently reported increase in metapopulation synchrony of the Glanville fritillary butterfly (Melitaea cinxia) in the Finnish archipelago, to see if it could be explained by an increase in synchrony of weather conditions. For this, we used 23 years of butterfly survey data together with monthly weather records for the same period. We first examined the associations between population growth rates within different regions of the metapopulation and weather conditions during different life‐history stages of the butterfly. We then examined the association between the trends in the synchrony of the weather conditions and the synchrony of the butterfly metapopulation dynamics. We found that precipitation from spring to late summer are associated with the M. cinxia per capita growth rate, with early summer conditions being most important. We further found that the increase in metapopulation synchrony is paralleled by an increase in the synchrony of weather conditions. Alternative explanations for spatial synchrony, such as increased dispersal or trophic interactions with a specialist parasitoid, did not show paralleled trends and are not supported. The climate driven increase in M. cinxia metapopulation synchrony suggests that climate change can increase extinction risk of spatially structured populations living in fragmented landscapes by altering their dynamics.     

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.     

Metapopulation dynamics of the bog fritillary butterfly: modelling the effect of habitat fragmentation

Population viability analysis (PVA) and metapopulation theory are valuable tools to model the dynamics of spatially structured populations. In this article we used a spatially realistic population dynamic model to simulate the trajectory of a Proclossiana eunomia metapopulation in a network of habitat patches located in the Belgian Ardenne. Sensitivity analysis was used to evaluate the relative influence of the different parameters on the model output. We simulated habitat loss by removing a percentage of the original habitat, proportionally in each habitat patch. Additionally, we evaluated isolation and fragmentation effects by removing and dividing habitat patches from the network, respectively. The model predicted a slow decline of the metapopulation size and occupancy. Extinction risks predicted by the model were highly sensitive to environmental stochasticity and carrying capacity. For a determined level of habitat destruction, the expected lifetime of the metapopulation was highly dependent on the spatial configuration of the landscape. Moreover, when the proportion of removed habitat is above 40% of the original habitat, the loss of whole patches invariably leads to the strongest reduction in metapopulation viability.     

Toward ecologically scaled landscape indices

Nature conservation is increasingly based on a landscape approach rather than a species approach. Landscape planning that includes nature conservation goals requires integrated ecological tools. However, species differ widely in their response to landscape change. We propose a framework of ecologically scaled landscape indices that takes into account this variation. Our approach is based on a combination of field studies of spatially structured populations (metapopulations) and model simulations in artificial landscapes. From these, we seek generalities in the relationship among species features, landscape indices, and metapopulation viability. The concept of ecological species profiles is used to group species according to characteristics that are important in metapopulations' response to landscape change: individual area requirements as the dominant characteristic of extinction risk in landscape patches and dispersal distance as the main determinant of the ability to colonize patches. The ecological profiles and landscape indices are then integrated into two ecologically scaled landscape indices (ESLI): average patch carrying capacity and average patch connectivity. The field data show that the fraction of occupied habitat patches is correlated with the two ESLI. To put the ESLI into a perspective of metapopulation persistence, we determine the viability for six ecological profiles at different degrees of habitat fragmentation using a metapopulation model and computer-generated landscapes. The model results show that the fraction of occupied patches is a good indicator for metapopulation viability. We discuss how ecological profiles, ESLI, and the viability threshold can be applied for landscape planning and design in nature conservation.     

Using connectivity metrics in conservation planning – when does habitat quality matter?

Aim The objective of conservation planning is often to prioritize patches based on their estimated contribution to metapopulation or metacommunity viability. The contribution that an individual patch makes will depend on its intrinsic characteristics, such as habitat quality, as well as its location relative to other patches, its connectivity. Here we systematically evaluate five patch value metrics to determine the importance of including an estimate of habitat quality into the metrics. Location We tested the metrics in landscapes designed to represent different degrees of variability in patch quality and different levels of patch aggregation. Methods In each landscape, we simulated population dynamics using a spatially explicit, continuous time metapopulation model linked to within patch logistic growth models. We tested five metrics that are used to estimate the contribution that a patch makes to metapopulation viability: two versions of the probability of connectivity index, two versions of patch centrality (a graph theory metric) and the metapopulation capacity metric. Results All metrics performed best in environments where patch quality was very variable and high quality patches were aggregated. Metrics that incorporated some measure of patch quality did better in all environments, but did particularly well in environments with high variance of patch quality and spatial aggregation of good quality patches. Main conclusions Including an estimate of patch quality significantly increased the ability of a given connectivity metric to rank correctly habitat patches according to their contribution to metapopulation viability. Incorporating patch quality is particularly important in landscapes where habitat quality is highly variable and good quality patches are spatially aggregated. However, caution should be used when applying patch metrics to homogeneous landscapes, even if good estimates of patch quality are available. Our results demonstrate that landscape structure and the degree of variability in patch quality need to be assessed prior to selecting a suitable method for estimating patch value.     

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.     

Drivers of local densities of endangered Lycaena helle butterflies in a fragmented landscape

Due to their specialised habitat requirements, butterflies are particularly vulnerable to habitat loss and fragmentation. Understanding the drivers of local abundances of species is essential for their effective conservation in fragmented landscapes. We investigated factors affecting population densities of an endangered European butterfly, the Violet Copper (Lycaena helle), occurring in a small metapopulation near the city of Kraków, southern Poland. The environmental parameters tested as predictors of the local densities of the species included both the variables associated with spatial structure of habitats such as patch sizes, their isolation and fragmentation as well as those potentially reflecting habitat quality. Patch area and vegetation height turned out to be the only factors significantly influencing L. helle densities, both having a positive effect. The positive impact of patch area is a bit surprising, since its relationship with population densities is typically negative in butterflies. In our study system it is likely to derive from source-sink dynamics as the smaller habitat patches are apparently too small to sustain viable local populations. In turn, the positive influence of vegetation height implies that the ongoing succession does not deteriorate the quality of the recently abandoned meadows yet, whereas higher turf may provide better sheltering places. The loss of almost half of L. helle habitat patches in the study area in recent years is alarming. However, its inclusion into the Natura 2000 system should help to conserve the species as long as this act is followed by proper management of its habitats.     

Synchrony in population counts predicts butterfly movement frequencies

1. Measuring functional connectivity, the ability of species to move between resource patches, is essential for conservation in fragmented landscapes. However, current methods are highly time consuming and expensive. 2. Population synchrony‐ the correlation in time series of counts between two long‐term population monitoring sites, has been suggested as a possible proxy measure of functional connectivity. To date, population synchrony has been shown to correlate with proxies for movement frequency such as the coverage of suitable habitat types in intervening landscapes, and also least cost distances around hostile land cover types. 3. This provides tentative evidence that synchrony is directly driven by movements of the focal species, but an alternative explanation is that these land cover types affect the movement of interacting species (e.g. natural enemies of the focal species) which can also drive synchronous population dynamics. Therefore, what is needed is a test directly relating population synchrony to movement frequencies of a focal species. 4. Here we use data from a 21 year mark‐release‐recapture study and show that population synchrony does indeed predict movements of a focal butterfly species Boloria eunomia (Esper). 5. There is growing evidence that population synchrony can be a useful conservation tool to measure functional connectivity.     

Patch occupancy and abundance of local populations in landscapes differing in degree of habitat fragmentation: a case study of the colonial black‐headed gull,Chroicocephalus ridibundus

Aim This study investigated whether habitat fragmentation at the landscape level influences patch occupancy and abundance of the black‐headed gull, Chroicocephalus ridibundus, and whether the response of the species to environmental factors is consistent across replicated landscape plots. Location Water bodies (habitat patches) in southern Poland. Methods Surveys were conducted in two landscape types (four plots in each): (1) more‐fragmented landscape, in which habitat patches were small (mean size 2.2–6.2 ha) and far apart (mean distance 2.5–3.1 km); and (2) less‐fragmented landscape, in which habitat patches were large (mean size 9.2–16.5 ha) and separated by short distances (mean 0.9–1.4 km). Observations were performed twice in 284 potential habitat patches during the 2007 breeding season. Results Colonies were significantly more frequent and larger in the less‐fragmented landscapes than in the more‐fragmented ones. Probability of patch occupancy and number of breeding birds were positively related with patch size and these relationships were especially strong in the more‐fragmented landscapes. In the less‐fragmented landscapes, the occurrence of black‐headed gulls was negatively related to the distance to the nearest local population, but in the more‐fragmented landscapes such a relationship was not detected. As distance to the nearest habitat patch increased, the probability of the patch occupancy decreased in the more‐fragmented landscapes. Moreover, abundance was negatively influenced by distance to the nearest habitat patch, especially strongly in more‐fragmented landscapes. Proximity of corridors (rivers) positively influenced the occupation of patches regardless of landscape type. The number of islets positively influenced occupancy and abundance of local populations, and this relationship was stronger in the more‐fragmented landscapes. Main conclusions Our results are in agreement with predictions from metapopulation theory and are the first evidence that populations of black‐headed gulls may have a metapopulation structure. However, patch occupancy and abundance were differentially affected by explanatory variables in the more‐fragmented landscapes than in the less‐fragmented ones. This implies that it is impossible to derive, a priori, predictions about presence/abundance patterns based on only a single landscape.     

Viability analyses with habitat-based metapopulation models

Population viability analysis (PVA) models incorporate spatial dynamics in different ways. At one extreme are the occupancy models that are based on the number of occupied populations. The simplest occupancy models ignore the location of populations. At the other extreme are individual-based models, which describe the spatial structure with the location of each individual in the population, or the location of territories or home ranges. In between these are spatially structured metapopulation models that describe the dynamics of each population with structured demographic models and incorporate spatial dynamics by modeling dispersal and temporal correlation among populations. Both dispersal and correlation between each pair of populations depend on the location of the populations, making these models spatially structured. In this article, I describe a method that expands spatially structured metapopulation models by incorporating information about habitat relationships of the species and the characteristics of the landscape in which the metapopulation exists. This method uses a habitat suitability map to determine the spatial structure of the metapopulation, including the number, size, and location of habitat patches in which subpopulations of the metapopulation live. The habitat suitability map can be calculated in a number of different ways, including statistical analyses (such as logistic regression) that find the relationship between the occurrence (or, density) of the species and independent variables which describe its habitat requirements. The habitat suitability map is then used to calculate the spatial structure of the metapopulation, based on species-specific characteristics such as the home range size, dispersal distance, and minimum habitat suitability for reproduction. Received: April 1, 1999 / Accepted: October 29, 1999     

Long-term demographic surveys reveal a consistent relationship between average occupancy and abundance within local populations of a butterfly metapopulation

Species distribution models are the tool of choice for large-scale population monitoring, environmental association studies and predictions of range shifts under future environmental conditions. Available data and familiarity of the tools rather than the underlying population dynamics often dictate the choice of specific method – especially for the case of presence–absence data. Yet, for predictive purposes, the relationship between occupancy and abundance embodied in the models should reflect the actual population dynamics of the modelled species. To understand the relationship of occupancy and abundance in a heterogeneous landscape at the scale of local populations, we built a spatio-temporal regression model of populations of the Glanville fritillary butterfly Melitaea cinxia in a Baltic Sea archipelago. Our data comprised nineteen years of habitat surveys and snapshot data of land use in the region. We used variance partitioning to quantify relative contributions of land use, habitat quality and metapopulation covariates. The model revealed a consistent and positive, but noisy relationship between average occupancy and mean abundance in local populations. Patterns of abundance were highly variable across years, with large uncorrelated random variation and strong local population stochasticity. In contrast, the spatio-temporal random effect, habitat quality, population connectivity and patch size explained variation in occupancy, vindicating metapopulation theory as the basis for modelling occupancy patterns in fragmented landscapes. Previous abundance was an important predictor in the occupancy model, which points to a spillover of abundance into occupancy dynamics. While occupancy models can successfully model large-scale population structure and average occupancy, extinction probability estimates for local populations derived from occupancy-only models are overconfident, as extinction risk is dependent on actual, not average, abundance.     

Potential metapopulation structure and the effects of habitat quality on population size of the endangered False Ringlet butterfly

The False Ringlet (Coenonympha oedippus) is a European butterfly species, endangered due to the severe loss and fragmentation of its habitat. In Hungary, two remaining populations of the butterfly occur in lowland Purple Moorgrass meadows. We studied a metapopulation occupying twelve habitat patches in Central Hungary. Our aim was to reveal what measures of habitat quality affect population size and density of this metapopulation, estimate dispersal parameters and describe phenology of subpopulations. Local population sizes and dispersal parameters were estimated from an extensive mark–release–recapture dataset, while habitat quality was characterized by groundwater level, cover of grass tussocks, bush cover, height of vegetation and grass litter at each habitat patch. The estimated size of the metapopulation was more than 3,000 individuals. We estimated a low dispersal capacity, especially for females, indicating a very low probability of (re)colonization. Butterfly abundance and density in local populations increased with higher grass litter, lower groundwater level and larger area covered by tussocks. We suppose that these environmental factors affect butterfly abundance by determining the microclimatic conditions for both larvae and adult butterflies. Our results suggest that the long-term preservation of the studied metapopulation needs the maintenance of high quality habitat patches by appropriate mowing regime and water regulation. Management also should facilitate dispersal to strengthen metapopulation structure with creating stepping-stones or gradually increase habitat quality in present matrix.     

Biased correlated random walk and foray loop: which movement hypothesis drives a butterfly metapopulation?

Animals in fragmented landscapes have a major challenge to move between high-quality habitat patches through lower-quality matrix. Two current mechanistic hypotheses that describe the movement used by animals outside of their preferred patches (e.g., high-quality habitat or home range) are the biased, correlated random walk (BCRW) and the foray loop (FL). There is also a variant of FL with directed movement (FL_dm). While these have been most extensively tested on butterflies, they have never been tested simultaneously with data across a whole metapopulation and over multiple generations, two key scales for population dynamics. Using the pattern-oriented approach, we compare support for these competing hypotheses with a spatially explicit individual-based simulation model on an 11-year dataset that follows 12 patches of the federally endangered Fender’s blue butterfly (Plebejus icarioides fenderi) in Oregon’s Willamette Valley. BCRW and medium-scale FL and FL_dm scenarios predicted the annual total metapopulation size for ≥9 of 12 patches as well as patch extinctions. The key difference, however, was that the FL scenarios predicted patch colonizations and persistence poorly, failing to adequately capture movement dynamics; BCRW and one FL_dm scenario predicted the observed patch colonization and persistence with reasonable probabilities. This one FL_dm scenario, however, had larger prediction intervals. BCRW, the biologically simplest and thus most parsimonious movement hypothesis, performed consistently well across all nine different tests, resulting in the highest quality metapopulation predictions for butterfly conservation.     

Modelling mortality and dispersal: consequences of parameter generalisation on metapopulation dynamics

Modelling dispersal is a fundamental step in the design of population viability analyses. Here, we address the question of the generalisation of population viability analysis models across landscapes by comparing dispersal between two metapopulations of the bog fritillary butterfly ( Proclossiana eunomia ) living in similar highly fragmented landscapes (<1% of suitable habitat in 9 km²). Differences in dispersal patterns were investigated using the virtual migration (VM) model, which was parameterised with capture–mark–recapture data collected during several years in both landscapes. The VM model allows the estimation of 6 parameters describing dispersal and mortality as well as the simulation of dispersal in the landscapes. The model revealed large differences in the VM parameter estimates between the two landscapes and consequently, simulations indicated differential rates of emigration and dispersal mortality. Furthermore, results from crossed-simulations i.e. simulations performed in one of the landscape but using parameter estimates from the other landscape emphasize that dispersal parameters are very specific to each metapopulation and to their landscape. Hence, we urge conservation biologists to be cautious with such parameter generalisations, even for the same species in comparable landscapes.     

Using surrogate data in population viability analysis: the case of the critically endangered cranberry fritillary butterfly

Population viability analyses (PVA) are central tools for the management of threatened populations. However, the parameterisation of effective PVA models is very demanding in high quality data, which are often impossible to collect on endangered populations. Here we propose the use of a generalisation strategy to bypass this limitation: management measures for an endangered metapopulation of the cranberry fritillary butterfly in the Netherlands are evaluated with RAMAS/GIS by using parameters estimated from a healthier metapopulation in Belgium. The Belgian metapopulation seems viable, with stable abundance and number of local populations, despite their erratic dynamics, whereas the Dutch metapopulation shows a continuous decline in the course of time, with many vacant habitat patches. Simulations of various scenarios indicated that (1) large scale restoration of habitat patches would be necessary to ensure long-term survival of the species in the Netherlands as not enough suitable habitats are currently remaining; and that (2) global warming is expected to put a major threat on both metapopulations by reducing the growth rate of this glacial relict species, and/or increasing environmental stochasticity (amplified climatic variations).     

Isolation determines patterns of species presence in highly fragmented landscapes

In fragmented landscapes, changes in habitat availability, patch size, shape and isolation may affect survival of local populations. Proposing efficient conservation strategies for such species relies initially on distinguishing the particular effects of those factors. To address these issues, we investigated the occurrence of 3 bird species in fragmented Brazilian Atlantic Forest landscapes. Playback techniques were used to collect presence/absence data of these species inside 80 forest patches, and incidence models were used to infer their occupancy pattern from landscape spatial structure. The relative importance of patch size, shape and surrounding forest cover and isolation was assessed using a model selection approach based on maximum likelihood estimation. The presence of all species was in general positively affected by the amount of surrounding habitat and negatively affected by inter‐patch distances. The joint effects of patch size and the surrounding landscape characteristics were important determinants of occupancy for two species. The third species was affected only by forest cover and mean patch isolation. Our results suggest that local species presence is in general more influenced by the isolation from surrounding forests than by patch size alone. We found evidence that, in highly fragmented landscapes, birds that can not find patches large enough to settle may be able to overcome short distances through the matrix and include several nearby patches within their home‐ranges to complement their resource needs. In these cases, patches must be defined as functionally connected habitat networks rather than mere continuous forest segments. Bird conservation strategies in the Atlantic forest should focus on increasing patch density and connectivity, in order to implement forest networks that reduce the functional isolation between large remnants with remaining core habitat.     

How Do Landscape Structure,Management and Habitat Quality Drive the Colonization of Habitat Patches by the Dryad Butterfly (Lepidoptera: Satyrinae) in Fragmented Grassland?

Most studies dealing with species distribution patterns on fragmented landscapes focus on the characteristics of habitat patches that influence local occurrence and abundance, but they tend to neglect the question of what drives colonization of previously unoccupied patches. In a study of the dryad butterfly, we combined classical approaches derived from metapopulation theory and landscape ecology to investigate the factors driving colonization from a recent refugium. In three consecutive transect surveys, we recorded the presence and numbers of imagos in 27 patches of xerothermic grassland and 26 patches of wet meadow. Among the predictors affecting the occurrence and abundance of the dryad, we considered environmental variables reflecting (i) habitat patch quality (e.g., goldenrod cover, shrub density, vegetation height); (ii) factors associated with habitat spatial structure (patch size, patch isolation and fragmentation); and (iii) features of patch surroundings (100-m buffers around patches) that potentially pose barriers or provide corridors. Patch colonization by the dryad was strongly limited by the distance from the species refugium in the region; there was a slight positive effect of shrub density in this respect. Butterfly abundance increased in smaller and more fragmented habitat patches; it was negatively impacted by invasive goldenrod cover, and positively influenced by the density of watercourses in patch surroundings. Nectar plant availability was positively related to species abundance in xerothermic grassland, while in wet meadow the effect was the reverse. We conclude that dryad colonization of our study area is very recent, since the most important factor limiting colonization was distance from the refugium, while the habitat quality of target patches had less relevance. In order to preserve the species, conservation managers should focus on enhancing the quality of large patches and should also direct their efforts on smaller and more fragmented ones, including those with relatively low resource availability, because such habitat fragments have an important role to play for specialist species.     

Modelling the effect of habitat fragmentation on range expansion in a butterfly

There is an increasing need for conservation programmes to make quantitative predictions of biodiversity responses to changed environments. Such predictions will be particularly important to promote species recovery in fragmented landscapes, and to understand and facilitate distribution responses to climate change. Here, we model expansion rates of a test species (a rare butterfly, Hesperia comma) in five landscapes over 18 years (generations), using a metapopulation model (the incidence function model). Expansion rates increased with the area, quality and proximity of habitat patches available for colonization, with predicted expansion rates closely matching observed rates in test landscapes. Habitat fragmentation constrained expansion, but in a predictable way, suggesting that it will prove feasible both to understand variation in expansion rates and to develop conservation programmes to increase rates of range expansion in such species.