The relative importance of patch habitat quality and landscape attributes on a declining steppe-bird metapopulation

Metapopulation theory is one of the most popular approaches to identify the factors affecting the spatial and temporal dynamics of populations in fragmented habitat networks. Habitat quality, patch area and isolation are mainly focused on when analyzing distribution patterns in fragmented landscapes. The effects of landscape heterogeneity in the non-occupied matrix, however, have been largely neglected. Here, we determined the relative importance of patch quality and landscape attributes on the occurrence, density and extinction of the Dupont’s lark (Chersophilus duponti), an endangered steppe passerine whose habitat has been extremely reduced to highly isolated and fragmented patches embedded in a mainly unsuitable landscape matrix. Habitat patch quality, measured in terms of vegetation structure, grazing pressure, arthropod availability, predator abundance, and inter-specific competition, did not affect occurrence, density or extinction. At the landscape scale, however, the species’ occurrence was principally determined by the interactions among patch size, geographic isolation and landscape matrix. Isolation had the main independent contribution to explaining the probability of occurrence, followed by landscape matrix composition and patch size. The species’ density was negatively correlated to patch size, suggesting crowding effects in small fragments, while extinction events were exclusively related to isolation. Our findings suggest that landscape rather than local population characteristics are crucial in determining the patterns of distribution and abundance of non-equilibrium populations in highly fragmented habitat networks. Consequently, conservation measures for these species should simultaneously involve patch size, isolation and landscape matrix and apply to the entire metapopulation rather than to particular patches.     

Spatial population structure of the Cabrera vole in Mediterranean farmland: The relative role of patch and matrix effects

There is mounting evidence that both patch networks and the intervening matrix influence species persistence in fragmented landscapes, though the relative importance of each of these factors in determining spatial population structure remains poorly understood. This study examined this issue using a three-year data set on the distribution of Cabrera voles (Microtus cabrerae) in Mediterranean farmland. The spatial pattern appeared consistent with a metapopulation structure, as voles occupied discrete tall herb patches scattered across the agricultural landscape, where local extinctions and colonizations induced temporal changes in occupancy patterns. Patch dynamics determined deviations from classical metapopulation assumptions, with over half the extinctions resulting from agricultural disturbance or vegetation succession, and recolonizations often occurring after the recovery of suitable habitat conditions sometime after disturbance. Occupancy in undisturbed patches was more stable, with vole occurrence in one year strongly reflecting that in the previous year. Overall, occupancy increased with both patch size and connectivity, but the unique contribution of patch variables to explain variation in vole occurrence was far smaller than that of matrix attributes. Voles occurred more often in patches surrounded by natural pastures, while prevalence declined with increasing cover by shrubland, pine plantations, improved pastures and grazed cropland. It is hypothesised that unfavourable land uses may increase the effective isolation of habitat patches through increased predation risk of dispersing voles. Conservation of the Cabrera vole in Mediterranean farmland should thus strive to maintain lightly grazed fields surrounding well-connected networks of suitable habitat patches.     

Metapopulation dynamics and conservation of the marsh fritillary butterfly: Population viability analysis and management options for a critically endangered species in Western Europe

This study investigates the dynamics and viability of a marsh fritillary butterfly Euphydryas aurinia metapopulation in a Belgian successional landscape. Based on capture-mark-recapture and winter nest census data, we first estimated demography (survival and recruitment rates, population size, density dependence) and dispersal parameters (emigration rate, effect of patch connectivity on dispersal, mortality during dispersal). Then using RAMAS/GIS platform, we parameterised a population viability analysis (PVA) model with these parameters to simulate the future of this metapopulation under different scenarios.The metapopulation does not seem viable even if natural reforestation is controlled by adequate management. In its present state, the patch system is not able to sustain enough individuals: due to the large temporal fluctuations in demographic parameters, a carrying capacity far higher than currently would be necessary to limit extinction risk to 1%, suggesting the existence of an extinction debt for the species in Belgium. The situation of E. aurinia appears much worse compared to two other fritillary species threatened in Belgium, for which similar PVA are available. It is therefore urgent to increase the carrying capacity of the patch system. How and where it is achieved are of secondary importance for the gain in viability: improvement of habitat quality through restoration, or increase of habitat quantity via enlargement of existing patches and/or creation of new habitat in the matrix. A regime of management based on regular re-opening and maintenance of habitat patches may be the only guarantee of long-term persistence for this critically endangered species in Belgium.     

Population fragmentation and extinction in the Iberian lynx

We studied the relationship between extinction frequencies of Iberian lynx subpopulations (Lynx pardinus) and their size and isolation during a 35-year period of strong geographic range contraction. At the end of this period there were fewer fragmentation events, fewer lynx populations of small size, and less isolation between them, than in simulated geographic ranges derived from a random distribution of local extinctions. Only small populations occupying <500 km² went extinct. Local extinction in large, self-sustainable populations probably resulted from the sole action of deterministic factors, e.g. widespread prey decline. As compared with large populations, small ones experienced increased contraction per unit occupied area, which may reflect demographic unstability. The consistent effect of isolation on extinction suggests that such unstability was often prompted by reduced immigration and the subsequent disruption of metapopulation equilibrium. Several practical recommendations could be derived. Provided that habitat quality was adequate, a lynx population should avoid extinction within 35 years if it occupied an area of at least 500 km². The persistence of small populations will also be enhanced by minimizing the distances to neighbouring populations within 30 km, and by maximizing the area occupied by these neighbours. Therefore, habitat management, or any other restoration action, directed to expand the area occupied by a small lynx population should be best located at points of its boundaries oriented towards other existing nearby populations.     

The role of fragment area and isolation in the conservation of heathland species

Heathland area in the northwestern part of Belgium has been strongly reduced during the past 200 years, and the remaining heathland is forced back into several small and isolated relics. In this study, we investigated how the fragmentation of these heathlands affects the distribution patterns of heathland plant species. Furthermore, we tested whether differences in patch occupancy patterns could be explained in terms of life history traits related to dispersal capacity and persistence. Multivariate logistic regression models showed that the incidence of almost three quarters of the species was influenced by fragmentation. For the majority of these species, isolation was the most important factor determining their presence or absence in a heathland patch. Differences between the species in isolation-sensitivity could be attributed mainly to differences in seed bank characteristics, with species having a long living seed bank being less affected by isolation. In contrast, species having mechanisms facilitating long distance dispersal were as much affected by isolation as species lacking these mechanisms.Our results suggest that for the majority of the species extinction in a patch can be prevented by dispersal from neighbouring patches. Further isolation of the patches should therefore be prevented and connectivity between the patches needs to be assured. As almost none of the species is affected by a declining patch area, for most even small patches are important for their survival. Hence, conservation efforts should focus not only on large heathlands.     

Viability and patch occupancy of a swamp rabbit metapopulation at the northern edge of its distribution

Swamp rabbits (Sylvilagus aquaticus) are state-endangered in Indiana, USA, and population decline has been attributed to habitat loss. We conducted pellet surveys as part of a long-term survey effort that has been conducted at approximate 10-year intervals over the last 40 years. We modeled patch occupancy and conducted a spatially-explicit population viability analysis (PVA). Although occupancy of individual patches varied over time, occupancy rate has been constant for the last 30 years, and Indiana swamp rabbits exist as a metapopulation that appears to be stable. Metapopulation dynamics were best characterized as being stationary, but area was an important factor in extinction rates; occupied patches (142 ± 37 ha) were significantly larger (P = 0.01) than unoccupied patches (79 ± 20 ha). We did not find strong support for models with colonization rates as a function of distance to neighboring patches, nor was distance to contiguous patches of habitat significantly different (P = 0.12) for occupied and unoccupied sites. Population viability analysis corroborated our findings based on occupancy modeling, and evaluation of the PVA model using occupancy data for the period 1985–2006 resulted in predictions that nearly matched our field observations (33% observed patch occupancy vs. 25% predicted patch occupancy). Population viability was most sensitive to reductions in survival and fecundity rates, but was otherwise robust to changes in parameters such as initial abundance and carrying capacity. Our findings provide novel insights into a poorly studied member of Sylvilagus and into species metapopulation dynamics at the edge of the range.     

Do epiphytic orchids behave as metapopulations? Evidence from colonization, extinction rates and asynchronous population dynamics

Previous in situ studies of orchid population dynamics with conservation relevance have focused on one or a few populations in a limited area. Many species of orchids occur as hyperdispersed populations in ephemeral habitats (epiphytic, twig epiphytes, short lived or vulnerable host). In this contribution, we show that orchid populations that are patchily distributed and that exist in disturbance-prone environments may act somewhat like a metapopulation with high turnover and low correlation in population dynamics. We tested for evidence of metapopulation dynamics in the riparian orchid Lepanthes rupestris by sampling over 1000 sites (250 initially occupied, 750 initially unoccupied) in biannual surveys for 5 years. Extinction and colonization of groups of orchids on a single substrate or patch (either trees or boulders) was common and more or less consistent across different time periods, and asynchronous subpopulation dynamics were evident among the populations. From this we predict non-zero equilibrium values for site occupancy () of L. rupestris. Nevertheless, this study species differed from a typical Levins’ metapopulation system in that small populations were more likely to go extinct than large populations, and that colonization of previously occupied sites was more common than colonization of initially unoccupied sites suggesting that site quality may influence population persistence and colonization. A major difficulty applying the metapopulation approach to orchid conservation is identifying empty sites suitable for colonization. In spite of this limitation, our study highlights the necessity of following multiple orchid subpopulations (e.g., an entire orchid “metapopulation” in the broad sense) may provide a more accurate basis for predicting persistence in epiphytic orchids.     

The effect of habitat fragmentation on communities of mutualists: Amazonian ants and their host plants

The consequences of fragmentation for communities of mutualist partners are for the most part unknown; moreover, most studies addressing this issue have been conducted on plant-pollinator communities. We evaluated how the experimental fragmentation of lowland Amazonian rain forest influenced a community of ant-plant mutualists. We inventoried a total of 1057 myrmecophytes in four fragments and four continuous forest sites; the twelve plant species recorded were occupied by 33 ant morphospecies, of which 11 were obligate plant inhabitants. Neither plant species richness, ant species richness, nor total ant-plant density were significantly lower in forest fragments. However, eight of the plant species, including three of the four most common, had higher mean densities in continuous forest than fragments. Of these four species, only one (Cordia nodosa) had significantly different colonization rates between habitat types, with higher colonization rates of plants in fragments. This may be because the Azteca species it is associated with increases in abundance in forest isolates. Although our results suggest that communities of ant-plant mutualists are likely to persist in fragmented tropical landscapes 25 years after fragment isolation, most species are rare and populations sizes in fragments are extremely low. Environmental and demographic stochasticity could therefore limit long-term population viability. We suggest future studies focus on evaluating how fragmentation has altered herbivore pressure and the dispersal of ants and plants to fragments, since the interaction of these factors is likely to have the greatest impact on long-term patterns of population persistence.     

Patch-occupancy models indicate human activity as major determinant of forest elephant Loxodonta cyclotis seasonal distribution in an industrial corridor in Gabon

The importance of human activity and ecological features in influencing African forest elephant ranging behaviour was investigated in the Rabi-Ndogo corridor of the Gamba Complex of Protected Areas in southwest Gabon. Locations in a wide geographical area with a range of environmental variables were selected for patch-occupancy surveys using elephant dung to assess seasonal presence and absence of elephants. Patch-occupancy procedures allowed for covariate modelling evaluating hypotheses for both occupancy in relation to human activity and ecological features, and detection probability in relation to vegetation density. The best fitting models for old and fresh dung data sets indicate that (1) detection probability for elephant dung is negatively related to the relative density of the vegetation, and (2) human activity, such as presence and infrastructure, are more closely associated with elephant distribution patterns than are ecological features, such as the presence of wetlands and preferred fresh fruit. Our findings emphasize the sensitivity of elephants to human disturbance, in this case infrastructure development associated with gas and oil production. Patch-occupancy methodology offers a viable alternative to current transect protocols for monitoring programs with multiple covariates.     

Probable consequences of high female mortality for speckled warblers living in habitat remnants

The effects of habitat fragmentation on the Australian avifauna have been widespread with species richness and abundance declining with reduced remnant size and habitat quality and increased habitat isolation. The speckled warbler, Chthonicola sagittata is one species from the highly fragmented temperate woodlands of eastern Australia that has declined across its range and populations that remain appear to be patchily distributed in habitat remnants. Specific causes of decline are unknown but several aspects of its biology make the species particularly vulnerable to decline in fragmented landscapes. Here, we analyse survey data (presence/absence) of speckled warblers in a large sample of habitat remnants from three regions to identify patterns of occupancy. We explore the effects of patch size on extinction risk using population viability analyses (PVA) and detailed demographic data from a behavioural study of individuals in the Australian Capital Territory, south-eastern Australia. Patch size was a strong predictor of the persistence of speckled warblers in habitat remnants. High density populations had higher probabilities of persisting, and inclusion of an Allee effect during drought decreased the probability of persistence. In the absence of an Allee effect, only high density populations in patches greater than 300 ha and low density populations in patches greater than700 ha had more than an 80% probability of persisting over 100 years. The accelerating decrease in population persistence below approximately 200-400 ha suggests that small populations were particularly vulnerable to stochastic demographic and environmental events. Adult female mortality was the single most important factor in driving population extinction. Our PVA model predictions matched the survey data for the Australian Capital Territory region remarkably well, but failed to predict occupancy rates in remnants in other regions. Differences in occupancy patterns between regions may, however, have resulted from inbreeding depression. This study demonstrates both the strengths and limitations of PVA analysis. PVA can predict occupancy patterns with reasonable accuracy, given good demographic data, but data for one region cannot be used universally for all regions. We highlight the need for studies of demography in different regions to interpret regional patterns of occupancy and to identify mechanisms of decline in remnant habitat.     

Persistence and conservation of a consumer-resource metapopulation with local overexploitation of resources

The monophagous flightless weevil Hadramphus spinipennis causes the frequent local extinction of its host plant Aciphylla dieffenbachii through overexploitation. Both species are endemic and restricted to the Chatham Islands group (New Zealand). The weevil is classified as endangered and the plant as a threatened species. As this herbivore-plant system is locally unstable long-term persistence only appears possible via a metapopulation structure in which subpopulations are connected by dispersal. This paper investigates this hypothesis using a spatially explicit model of a consumer population whose resource is patchily distributed. The parameters are adapted to the H. spinipennis-A. dieffenbachii system. Our model includes local population dynamics and dispersal of the consumer, the destruction of host plant patches due to foraging and their regeneration. The results show that the key factor for long-term persistence is the short-range dispersal of the consumer with high mortality during dispersal. Only this highly inefficient dispersal prevents the synchronisation of local dynamics while ensuring sufficient colonisation of regenerated patches. We also show that although synchronisation through spatially correlated environmental fluctuations may be critical for long-term persistence, it cannot replace the strong effects of dispersal. Thus, in a consumer-resource metapopulation with deterministic local extinction, the strength and spatial scale of consumer dispersal in relation to the spatial and temporal scales of the resource patch dynamics (patch destruction and regeneration, number, size and distance of patches) play a vital role for long-term persistence. The consequence for conservation management is that such metapopulations could in fact profit rather than suffer from decreasing connectivity of resource patches.     

Patch dynamics and the timing of colonization-abandonment events by male Kirtland’s Warblers in an early succession habitat

Habitat colonization and abandonment affects the distribution of a species in space and time, ultimately influencing the duration of time habitat is used and the total area of habitat occupied in any given year. Both aspects have important implications to long-term conservation planning. The importance of patch isolation and area to colonization-extinction events is well studied, but little information exists on how changing regional landscape structure and population dynamics influences the variability in the timing of patch colonization and abandonment events. We used 26 years of Kirtland’s Warbler (Dendroica kirtlandii) population data taken during a habitat restoration program (1979-2004) across its historical breeding range to examine the influence of patch attributes and temporal large-scale processes, specifically the rate of habitat turnover and fraction of occupied patches, on the year-to-year timing of patch colonization and abandonment since patch origin. We found the timing of patch colonization and abandonment was influenced by patch and large-scale regional factors. In this system, larger patches were typically colonized earlier (i.e., at a younger age) and abandoned later than smaller patches. Isolated patches (i.e., patches farther from another occupied patch) were generally colonized later and abandoned earlier. Patch habitat type affected colonization and abandonment; colonization occurred at similar patch ages between plantation and wildfire areas (9 and 8.5 years, respectively), but plantations were abandoned at earlier ages (13.9 years) than wildfire areas (16.4 years) resulting in shorter use. As the fraction of occupied patches increased, patches were colonized and abandoned at earlier ages. Patches were abandoned at older ages when the influx of new habitat patches was at low and high rates. Our results provide empirical support for the temporal influence of patch dynamics (i.e., patch destruction, creation, and succession) on local colonization and extinction processes that help explain large-scale patterns of habitat occupancy. Results highlight the need for practitioners to consider the timing of habitat restoration as well as total amount and spatial arrangement of habitat to sustain populations.     

Extirpation, colonization, and habitat dynamics of a keystone species along an urban gradient

Using 5 years of patch occupancy data for 384 habitat fragments, we evaluated population and habitat dynamics of the black-tailed prairie dog in urban habitat remnants in the rapidly developing landscape of Denver, CO, USA. Specifically, we evaluated the landscape factors, including fragment area, age, and connectivity, that characterize the habitat fragments most likely to be colonized by prairie dogs, as well as those experiencing local extinctions. In addition, we determined which patch types were most often removed by land development. Sites in proximity to colonies were more likely to be colonized by prairie dogs. Local extinctions were most common on isolated colonies, and older and more isolated colonies were more likely to be extirpated by human activity. In general, smaller and older habitat patches were at the highest risk of being lost to land development. Our results provide observations of dynamic changes to the distribution of a potential keystone species in an urban area, which can be used to inform island biogeographic and metapopulation models for wildlife persistence in developing landscapes. Although populations are currently in decline, most local extinctions are the direct result of human activity, and we suggest that prairie dogs in this area can persist with appropriate management.     

Agricultural landscape modification increases the abundance of an important food resource: Mistletoes, birds and brigalow

Agricultural landuse alters landscape pattern, with important consequences for native species. Complex responses to changes in landscape pattern may be observed for species with strong inter-specific relationships. We investigate how the abundance of an ecologically important food resource, the grey mistletoe (Amyema quandang), is influenced by the interactive influences of human-modified landscape pattern, bird disperser abundance and host condition. Mistletoe abundance was estimated in brigalow (Acacia harpophylla) dominated remnants in three separate study areas with differing landscape patterns. Negative binomial generalised linear models of mistletoe abundance were tested with combinations of seven explanatory variables: study area, landscape connectivity and edge contrast, patch shape, bird disperser abundance, brigalow foliage cover and basal area of standing dead trees. Ranking of model averaged parameter estimates showed that study area, bird disperser abundance, patch shape, dead trees and connectivity had the strongest influence on A. quandang abundance. Mistletoe abundance apparently increased with landscape modification, particularly in narrow linear patches, but may be limited by the availability of seed dispersers and host condition. Given high landscape modification and ongoing degradation, management should be targeted towards maintaining brigalow remnant condition to ensure bird and mistletoe populations can be supported in the long-term.     

Wild-captive metapopulation viability analysis

We developed an interactive management model for wild and captive populations of the ploughshare tortoise or angonoka, Geochelone yniphora. Interactive management is based on the translocation of individuals between wild and captive populations to simulate a metapopulation. Demographic parameters of one captive and two wild populations of this rare tortoise were used to conduct a metapopulation viability analysis (MVA). The effectiveness of the conservation strategy proposed for this species was then evaluated by modifying the probability of extinction and growth of the metapopulation over a fixed period of time. Several alternative scenarios of interactive management were then tested and ranked in terms of their effect on the viability of the metapopulation. The model predicted that catastrophic events such as bush fires would likely have a negative effect on the future of remaining wild populations. However, the model also predicted that the use of captive-born offspring to establish additional wild populations would decrease the risk of extinction of the metapopulation as a whole. We believe that, when supported by sound knowledge of the demographic parameters of a species, the use of MVA as part of an interactive management program can be an effective conservation tool that allows assessment of the probable response of both captive and wild populations to different management alternatives. One of the most interesting aspects of this interactive management approach is the link between in situ and ex situ conservation.     

Resource availability, matrix quality, microclimate, and spatial pattern as predictors of patch use by the Karner blue butterfly

Determination of which aspects of habitat quality and habitat spatial arrangement best account for variation in a species’ distribution can guide management for organisms such as the Karner blue butterfly (Lycaeides melissa samuelis), a federally endangered subspecies inhabiting savannas of Midwest and Eastern United States. We examined the extent to which three sets of predictors, (1) larval host plant (Lupinus perennis, wild lupine) availability, (2) characteristics of the matrix surrounding host plant patches, and (3) factors affecting a patch’s thermal environment, accounted for variation in lupine patch use by Karner blues at Indiana Dunes National Lakeshore, Indiana and Fort McCoy, Wisconsin, USA. Each predictor set accounted for 7-13% of variation in patch occupancy by Karner blues at both sites and in larval feeding activity among patches at Indiana Dunes. Patch area, an indicator of host plant availability, was an exception, accounting for 30% of variation in patch occupancy at Indiana Dunes. Spatially structured patterns of patch use across the landscape accounted for 9-16% of variation in patch use and explained more variation in larval feeding activity than did spatial autocorrelation between neighboring patches. Because of this broader spatial trend across sites, a given management action may be more effective in promoting patch use in some portions of the landscape than in others. Spatial trend, resource availability, matrix quality, and microclimate, in general, accounted for similar amounts of variation in patch use and each should be incorporated into habitat management planning for the Karner blue butterfly.     

Factors influencing the dispersion and fragmentation of endangered mountain caribou populations

Mountain caribou, an ecotype of woodland caribou, are endangered due to the loss and fragmentation of old forests on which they depend. However, a wider array of natural and human factors may limit caribou persistence and isolate populations, and understanding these may help to stop or reverse population declines by forecasting risk and targeting core habitat areas and key linkages for protection, enhancement or restoration. Across most of the historic range of mountain caribou, we conducted a bi-level analysis to evaluate factors related to the persistence of, and landscape occupancy within, remaining subpopulations. We used caribou location data from 235 radio-collared animals across 13 subpopulations to derive a landscape occupancy index, while accounting for inherent sampling biases. We analyzed this index against 33 landscape variables of forest overstory, land cover, terrain, climate, and human influence. At the metapopulation level, the persistence of subpopulations relative to historic range was explained by the extent of wet and very wet climatic conditions, the distribution of both old (>140 yr) forests, particularly of cedar and hemlock composition, and alpine areas. Other important factors were remoteness from human presence, low road density, and little motorized access. At the subpopulation level, the relative intensity of caribou landscape occupancy within subpopulation bounds was explained by the distribution of old cedar/hemlock and spruce/subalpine fir forests and the lack of deciduous forests. Other factors impeding population contiguity were icefields, non-forested alpine, hydro reservoirs, extensive road networks, and primary highway routes. Model outputs at both levels were combined to predict the potential for mountain caribou population persistence, isolation, and restoration. We combined this output with the original occupancy index to gauge the potential vulnerability of caribou to extirpation within landscapes known to have recently supported animals. We discuss implications as they pertain to range-wide caribou population connectivity and conservation.     

Classical metapopulation theory as a useful paradigm for the conservation of an endangered amphibian

Classical metapopulation theory (CMT) has proven an attractive paradigm for ecologists concerned with the conservation of aquatic-breeding amphibians, given its apparent fit with the population dynamics of these animals, and the opportunities the concept provides to assess alternate management options. Nevertheless, several authors have cautioned against uncritical application of this paradigm. We assessed the application of CMT to the conservation of the endangered growling grass frog (Litoria raniformis) in the urbanising landscapes of Melbourne, Victoria, Australia. Support for five predictions developed from the basic tenets of CMT was assessed using a multi-year occupancy and mark–recapture data-set. There was congruence between all five predictions and data. Wetland occupancy was strongly influenced by the proximity of neighbouring populations (‘connectivity’), but the estimated rate of dispersal between wetlands was low. Wetland occupancy was also temporally dynamic, with only a weak effect of connectivity on the probability of extinction, but a strong positive influence of connectivity on the probability of colonisation. Our work confirms that CMT provides a useful model of the dynamics of L. raniformis in urbanising landscapes, and justifies the application of the paradigm to conservation planning for this species. We argue that CMT may prove relevant to numerous aquatic-breeding amphibians, and encourage assessment of the application of CMT to the conservation of these animals.     

Factors affecting persistence of terrestrial orchids in wet meadows and implications for their conservation in a changing agricultural landscape

We examined the occurrence of Dactylorhiza majalis, the most abundant terrestrial orchid species growing in rapidly disappearing wet meadows, at 50 historical sites for 3 years. We aimed to find the most frequent reasons for its recent extinction at many sites. We found that the main reasons for its extinction were absence of mowing, intensive fertilisation and washouts of fertilisers from fields nearby. At extant sites, we studied its biometric characteristics and composition of surrounding vegetation, to determine factors affecting its persistence. Bad performance of persisting populations of this species was associated with prevalence of grasses, low May temperatures and absence of mowing. This confirms, at metapopulation level, what has previously been observed at the level of individual populations. We suggest that the system of agricultural subsidies in the country should change towards more sensitive allocation of funds to those farmers, who will adopt the appropriate management of wet meadows and their surroundings.