Effects of habitat fragmentation on the fitness of two common wetland species,Carex davalliana and Succisa pratensis

Small habitat size and spatial isolation may cause plant populations to suffer from genetic drift and inbreeding, leading to a reduced fitness of individual plants. We examined the germination, establishment, growth, and reproductive capacity of two characteristic species of mown fen meadows, Carex davalliana, and Succisa pratensis, common in Switzerland. Plants were grown from seeds, which were collected in 18 habitat islands, differing in size and in degree of isolation. We used both common garden and reciprocal transplant experiments to assess effects of habitat fragmentation. In the common garden, plants of Carex originating from small habitat islands yielded 35% less biomass, 30% fewer tillers, and 45% fewer flowering tillers than plants from larger ones. In contrast, plants of Succisa originating from small habitat islands yielded 19% more biomass, 14% more flower heads and 35% more flowers per flower head than plants from larger ones. Moreover, plants of Succisa from small isolated habitats yielded 32% more rosettes than did plants from small connected islands. Reciprocally transplanted plants of Succisa originating from small habitat islands produced 7% more rosettes than plants from larger ones. There was no effect of small habitat size and isolation on germination and establishment of both species in the field. Our results document genetic differences in performance attributable to habitat fragmentation in both species. We suggest that fitness loss in Carex is caused by inbreeding depression, whereas in Succisa the differences in fitness are more likely caused by genetic differentiation. Our study implies that habitat fragmentation affects common habitat-specific species, such as Carex and Succisa, as well as rare ones.     

Forest fragmentation and plant reproductive success: a case study in four perennial herbs

We assessed the effects of habitat fragmentation on reproductive success in natural populations of four forest herbs with differing life-history traits and whose distribution patterns appeared to be negatively affected by decreased habitat size and/or increased isolation: Carex sylvatica, Galium odoratum, Sanicula europaea and Veronica montana. Our aims were to test (1) whether habitat size and isolation are positively correlated with population size and isolation, respectively, (2) whether plant reproductive success, a major component of plant fitness, is reduced in small and/or isolated populations when also accounting for differences in habitat quality (edaphic conditions, light intensity) and the effects of plant size, and (3) whether species with different life histories are affected differently. There were significant positive relationships between habitat and population size and between habitat and population isolation in some, but not all of the species. We mostly found no negative effects of small population size or isolation on reproduction. However, reproductive success was reduced in small populations of Sanicula, and this effect was independent of differences in plant size and environmental conditions. The reduced fecundity in small populations may be a consequence of the Allee-effect, a possible mechanism being pollen limitation. Furthermore, the proportion of flowering ramets was reduced in small and isolated populations of Galium, which may have been caused by changes in population structure. Lastly, we found some evidence for largely outcrossing, non-clonal species to be more sensitive to reductions in population size, at least in terms of their reproductive success.     

Zonation of plant cover and environmental factors in wetlands of the Sanjiang Plain,northeast China

The zonation of depressional and riparian wetlands in the Sanjiang Plain of northeastern China was studied to describe their vegetation composition and environmental variables. We sampled 108 plots in 6 depression and riparian wetlands. Samples were classified into 4 groups using two‐way indicator species analysis (TWINSPAN). Emergent marsh vegetation was characterized by Carex lasiocarpa and C. pseudocuraica, meadow marsh vegetation by tussock species such as Carex appendiculata and C. meyeriana, wet meadow vegetation by Calamangrostis angustifolia and Carex appendiculata, and shrub meadow vegetation by the shrubs Betula fruticosa, Alnus sibirica and Salix rosmarinifolia and the graminids Carex schmidtii and Calamagrostis angustifolia. CCA ordination showed that water table, organic matter and available N were the major factors explaining the vegetation zonation pattern. Compared with other Northern Hemisphere regions, bog and fen vegetation are completely absent due to climatic conditions unfavorable for peat formation. Out of four vegetation types, only the Carex lasiocarpa community and the C. pseudo‐curaica community have been found in other regions. However, at the species level many species are widespread and some species are vicariant or pseudovicariant to other regions in the world. Our study suggests that topography and hydrology may be the most important determinants of the vegetation pattern in this region.     

Nutrient limitation in boreal rich‐fen vegetation: A fertilization experiment

Abstract. Rich‐fen vegetation influenced by hay‐making in the Sølendet Nature Reserve, Central Norway, was fertilized with N, P and K in a full‐factorial fertilization experiment to investigate the nutrient limitation of plant growth at both community and species levels. Above‐ground biomass, shoot density and nutrient concentration were measured in several species and groups of species at three sites after two years of fertilization. At the community level, the results indicate multiple limitation by N and P in the two least productive rich‐fen communities: one characterized by small sedges and herbs, and the other by high abundance of Menyanthes trifoliata and tall sedges. Increased nutrient availability had no effect on a more highly productive, tall‐growing, spring‐influenced community, indicating no nutrient limitation. The results at the species level correspond well with those at the community level, indicating multiple limitation by N and P in most of the dominant and sub‐dominant species. However, P seems to limit growth more than N in Succisa pratensis, and N seems to limit growth more than P in Carex panicea. Furthermore, Eriophorum angustifolium seems to be limited by K. The results did not show which nutrient limits the growth of Carex dioica, C. lasiocarpa and Trichophorum cespitosum. Indications that growth in low‐productive, boreal rich‐fen communities is generally limited by P was not confirmed.     

Key variables controlling the vegetation of a cool-temperate mire in northern Japan

Abstract. In the cool-temperate Bibi Mire, Hokkaido, Japan, valley fens and flood-plain fens have quite different vegetation. The main variables controlling the vegetation were all hydrological: mean water level, water level fluctuation and surface water flow. Chemical factors such as electrical conductivity, dissolved oxygen and related peat decomposition were less important. The pH was about neutral and has little effect. The flood-plain fen developed under fluctuating water table conditions. The dominant species are Calamagrostis langsdotffii and Carex pseudocuraica. When temporal inundation occurs in the rainy or typhoon seasons, the submergence stimulates bud germination of the stoloniferous C. pseudocuraica, which can rapidly elongate its stolons upward. Some large floating peat mats occurred in the flood-plain fen zone. On these mats some Alnus japonica saplings establish and patches of alder forest can arise. Here the water level was higher than in the peripheral alder forest zone. The valley fen is dominated by Carex lasiocarpa var. occultans and/or C. limosa. It is formed under stable water table conditions in the inundated parts of the mire -where the non-inundated wet areas are dominated by alder trees. In the area where the surface water is flowing, these two fen sedges grow in deeper water since the high oxygen content is considered to compensate the flooding stress.     

Effects of Small Habitat Size and Isolation on the Population Structure of Common Wetland Species

Abstract: Habitat fragmentation may cause plant species to suffer from inbreeding and genetic drift, which affects population viability negatively. Less viable populations may contain an altered population structure, i.e., they have a smaller proportion of seedlings and a larger proportion of vegetative adults, when compared to large extensive populations. We applied a hierarchical-spatial field design, consisting of large habitat islands with surrounding near and distant small habitat islands to distinguish between the two components of habitat fragmentation, i.e., the effect of small habitat size and the effect of isolation. We studied two common habitat-specific species: Carex davalliana (Cyperaceae) and Succisa pratensis (Dipsacaceae). We identified a decrease in the proportion of C. davalliana seedlings and an increase in the proportion of vegetative tillers in response to isolation. In S. pratensis, we identified a decrease in the proportion of seedlings and an increase in the proportion of vegetative rosettes that were attributable to small habitat size. Hence, we confirmed alterations in the population structure of those common species, although both species were affected by a different component of habitat fragmentation. The observed discrepancy between species could be related their life history. We conclude that habitat fragmentation adversely affects not only rare but also common species. Their loss of viability can inevitably lead to alterations in vegetation structure and consequently lead to the further loss of this species-rich and threatened ecosystem type.     

Comparisons of nutrient recovery and specific leaf area variation betweenCarex lasiocarpa var.occultans andCarex thunbergii var.appendiculata with reference to nutrient conditions and shading byPhragmites australis

The distribution of two sedge species was studied in two mires which differ in abiotic environments and in distribution ofPhragmites australis. Carex lasiocarpa var.occultans dominated in nutrient-poor valley mire, andCarex thunbergii var.appendiculata dominated in nutrient-rich flood plain subject to water fluctuations.Phragmites australis grew well in nutrient-rich conditions. The distribution ofC. lasiocarpa showed a strong negative correlation withP. australis coverage, whereasC. thunbergii coverage was not affected byP. australis. The leaf area per dry leaf mass (specific leaf area: SLA) ofC. thunbergii increased with shading byP. australis, but that ofC. lasiocarpa was stable. The SLA flexibility ofC. thunbergii to light interception might enable this species to invadeP. australis patches in nutrient-rich environments. The residual nutrient ratio of nitrogen and phosphorus (the ratio of the residual nutrient content at the end of the growing season to peak nutrient content) in the vegetative ramet ofC. thunbergii was 1.7 times higher than that ofC. lasiocarpa. This low residual ratio may indicate effective nutrient recovery to storage organs. The effective nutrient recovery inC. lasiocarpa might enable this species to grow even in nutrient-poor environments. However, it may be difficult forC. lasiocarpa to expand its habitat to nutrient-rich areas whereP. australis dominates as it is not shade tolerant.     

Vegetation and microclimatic edge effects in two mixed-mesophytic forest fragments

Forest edges are known to consist of microenvironments that may provide habitat for a different suite of species than forest interiors. Several abiotic attributes of the microenvironment may contribute to this change across the edge to center gradient (e.g., light, air temperature, soil moisture, humidity). Biotic components, such as seed dispersal, may also give rise to changes in species composition from forest edge to interior. We predicted that abiotic and biotic measures would correlate with distance from forest edge and would differ among aspects. To test these predictions, we measured abiotic and biotic variables on twelve 175 m transects in each of two 24 ha forest fragments in east-central Illinois that have remained in continuous isolation for upwards of 100 years. Both univariate and multivariate techniques were used to best describe the complex relationships among abiotic factors and between abiotic and biotic factors. Results indicate that microclimatic variables differ in the degree to and distance over which they show an edge effect. Relative humidity shows the widest edge, while light and soil moisture have the steepest gradients. Aspect influences are evidenced by the existence of more pronounced edge effects on south and west edges, except when these edges are protected by adjacent habitat. Edges bordered by agricultural fields have more extreme changes in microclimate than those bordered by trees. According to PCA results, species richness correlates well with microclimatic variation, especially light and soil moisture; however, in many cases species richness had a different depth of edge influence than either of these variables. The herbaceous plant community is heavily dominated by three species. Distributions of individual species as well as changes in plant community composition, estimated with a similarity index, indicate that competition may be influencing the response of the vegetation to the edge to interior gradient. This study indicates that edge effects must be considered when the size and potential buffering habitat of forest preserves are planned.     

Response of Plant Height,Species Richness and Aboveground Biomass to Flooding Gradient along Vegetation Zones in Floodplain Wetlands,Northeast China

Flooding regime changes resulting from natural and human activity have been projected to affect wetland plant community structures and functions. It is therefore important to conduct investigations across a range of flooding gradients to assess the impact of flooding depth on wetland vegetation. We conducted this study to identify the pattern of plant height, species richness and aboveground biomass variation along the flooding gradient in floodplain wetlands located in Northeast China. We found that the response of dominant species height to the flooding gradient depends on specific species, i.e., a quadratic response for Carex lasiocarpa, a negative correlation for Calamagrostis angustifolia, and no response for Carex appendiculata. Species richness showed an intermediate effect along the vegetation zone from marsh to wet meadow while aboveground biomass increased. When the communities were analysed separately, only the water table depth had significant impact on species richness for two Carex communities and no variable for C. angustifolia community, while height of dominant species influenced aboveground biomass. When the three above-mentioned communities were grouped together, variations in species richness were mainly determined by community type, water table depth and community mean height, while variations in aboveground biomass were driven by community type and the height of dominant species. These findings indicate that if habitat drying of these herbaceous wetlands in this region continues, then two Carex marshes would be replaced gradually by C. angustifolia wet meadow in the near future. This will lead to a reduction in biodiversity and an increase in productivity and carbon budget. Meanwhile, functional traits must be considered, and should be a focus of attention in future studies on the species diversity and ecosystem function in this region.     

Patch occupancy by stone martens Martes foina in fragmented landscapes of central Spain: the role of fragment size,isolation and habitat structure

We studied the response to forest fragmentation of a generalist carnivore, the stone marten Martes foina, in highly fragmented landscapes of central Spain. Five different areas (n = 178 fragments) in central Spain were surveyed. This paper analyses the relationship between fragment use by martens (measured through scat presence) and a series of variables related to the size, isolation and vegetation structure of each fragment by means of stepwise logistic regression. Size and isolation have an important effect on stone marten presence in fragments. Our results were similar to those found for other marten species in landscapes with coarse-grain fragmentation, but they contrast with other studies conducted in landscapes with fine-grain fragmentation. These data suggested that in highly fragmented landscapes, size and isolation factors resulting from forest fragmentation were responsible for determining marten responses, irrespective of their habitat generalism. Management policies for the stone marten in highly fragmented scenarios require the maintenance of large forests near continuous forest tracts in mountains or riparian woodlands.     

Potential effects of warming and drying on peatland plant community composition

Boreal peatlands may be particularly vulnerable to climate change, because temperature regimes that currently constrain biological activity in these regions are predicted to increase substantially within the next century. Changes in peatland plant community composition in response to climate change may alter nutrient availability, energy budgets, trace gas fluxes, and carbon storage. We investigated plant community response to warming and drying in a field mesocosm experiment in northern Minnesota, USA. Large intact soil monoliths removed from a bog and a fen received three infrared warming treatments crossed with three water‐table treatments (n = 3) for five years. Foliar cover of each species was estimated annually. In the bog, increases in soil temperature and decreases in water‐table elevation increased cover of shrubs by 50% and decreased cover of graminoids by 50%. The response of shrubs to warming was distinctly species‐specific, and ranged from increases (for Andromeda glaucophylla) to decreases (for Kalmia polifolia). In the fens, changes in plant cover were driven primarily by changes in water‐table elevation, and responses were species‐ and lifeform‐specific: increases in water‐table elevation increased cover of graminoids – in particular Carex lasiocarpa and Carex livida– as well as mosses. In contrast, decreases in water‐table elevation increased cover of shrubs, in particular A. glaucophylla and Chamaedaphne calyculata. The differential and sometimes opposite response of species and lifeforms to the treatments suggest that the structure and function of both bog and fen plant communities will change – in different directions or at different magnitudes – in response to warming and/or changes in water‐table elevation that may accompany regional or global climate change.     

The role of patch area and habitat diversity in explaining native plant species richness in disturbed suburban forest patches in northern Belgium

Relatively easy measurable patch characteristics (especially habitat diversity measures) have proven to be valuable indicators of forest plant species richness in forest fragments of relatively undisturbed areas. Urban and suburban forest patches, however, are characterized by a specific landscape ecological context implying that specific processes may influence ecosystem functioning and hence that other abiotic indicators for plant diversity are more appropriate. We studied the relation between functional ecological plant species groups and suburban forest patch characteristics such as patch area, habitat diversity and isolation. Some components of species richness were related to the isolation of the patches. In contrast to previous similar large-scale fragmentation studies in more rural areas, further results stressed the overwhelming importance of patch area relative to habitat variables in determining species richness. This suggests (1) the occurrence of density-dependent species extinction processes in small forest patches; or (2) the existence of external deterministic factors which put a major constraint on species richness in small patches. We tend to support the latter hypothesis and propose forest disturbance and associated black cherry (Prunus serotina Ehrh.) invasion as such a possible external factor. Small forest patches may be more sensitive to disturbance and biological invasion due to various reasons. Hence large forest patches are to be preferred for plant conservation in the suburban area.     

Richness and composition of plants and birds on land‐bridge islands: effects of island attributes and differential responses of species groups

Aim To test relationships between the richness and composition of vascular plants and birds and attributes of habitat fragments using a model land‐bridge island system, and to investigate whether the effects of fragmentation differ depending on species natural history traits. Location Thousand Island Lake, China. Methods We compiled presence/absence data of vascular plant and bird species through exhaustive surveys of 41 islands. Plant species were assigned to two categories: shade‐intolerant and shade‐tolerant species; bird species were assigned to three categories: edge, interior, and generalist species. We analysed the relationships between island attributes (area, isolation, elevation, shape complexity, and perimeter to area ratio) and species richness using generalized linear models (GLMs). We also investigated patterns of composition in relation to island attributes using ordination (redundancy analysis). Results We found that island area explained a high degree of variation in the species richness of all species groups. The slope of the species–area relationship (z) was 0.16 for all plant species and 0.11 for all bird species. The lowest z‐value was for generalist birds (0.04). The species richness of the three plant species groups was associated with island area per se, while that of all, generalist, and interior birds was explained mainly by elevation, and that of edge bird species was associated primarily with island shape. Patterns of species composition were most strongly related to elevation, island shape complexity, and perimeter to area ratio rather than to island area per se. Species richness had no significant relationship with isolation, but species composition did. We also found differential responses among the species groups to changes in island attributes. Main conclusions Within the Thousand Island Lake system, the effects of fragmentation on both bird and plant species appear to be scale‐dependent and taxon‐specific. The number of plant species occurring on an island is strongly correlated with island area, and the richness of birds and the species composition of plants and birds are associated with variables related to habitat heterogeneity. We conclude that the effects of fragmentation on species diversity and composition depend not only on the degree of habitat loss but also on the specific patterns of habitat fragmentation.     

Nutrient limitation in species-rich lowland fens

Abstract. Nitrogen, phosphorus and potassium were supplied to some Belgian fens of varying nutrient status and productivity. Plant growth in the lowest productive fen with a species-rich Caricion davallianae vegetation was strongly P-limited. N was ineffective when applied alone, but increased the effect of P-addition when applied together. Summer biomass and plant nutrient concentrations were monitored for four years, and showed partial recovery of nutrient limitation. In a more productive fen dominated by Carex lasiocarpa and in a fen meadow, nutrient limitation was less strong. N limited growth in the productive fen, and N and K were co-limiting in the fen meadow. The P-concentration in the productive fen vegetation showed a marked increase after P-fertilization, but it did not result in higher standing crop. The significance of P-limitation for the conservation of species rich low productive fens is discussed. P-limitation may be an essential feature in the conservation of low productive rich fens: because it is less mobile in the landscape than N and/or because it is an intrinsic property of this vegetation type. Plant nutrient concentrations and N:P-ratios may be used as an indication for the presence and type of nutrient limitation in the vegetation. We found N:P-ratios of 23 to 31 for a P-limited site and 8 to 15 in N-limited sites. This was in agreement with critical values from the literature: N:P > ca. 20 for P-limitation and N:P < 14 for N-limitation. Thus, this technique appears valid in the vegetation types that were studied here.     

Topographic position and water chemistry of fens in a Dutch river plain

Abstract. On the Vecht river plain (western Netherlands), small fens, remnants of a large mesotrophic wetland bordering a moraine, of 1 to 5 ha are found in a man-made matrix of lakes and pastures. The regional position of the fens, local position of sampling sites, composition of the vegetation and local hydrological variables were measured. Polders in the river plain produce a complex hydrology obscuring the regional zonation between moraine and river. Water supply and species composition are determined more by a site's regional than local position. High-productivity reedlands are abundant close to the river. Carex paniculata reedlands receive large amounts of river water, which gives their fen water a high K⁺ concentration. Low-productivity C. diandra fens and litter fens have their optimum closer to the moraine. C. diandra fens are fed mainly by inflowing nutrient-poor ground- or surface water; litter fens receive primarily rainwater. Nutrients in fen water and in peat are lowest in C. diandra and C. lasiocarpa fens, but do not differ significantly between the communities. In both, iron seems to be more important than calcium in reducing phosphate solubility. Iron richness in the C. diandra fens is caused by present inflows of ground- or surface water, while in C. lasiocarpa fens, which succeed the former, iron richness is the result of historical inflows.     

Edge,area and anthropization effects on mangrove-dwelling ant communities

In mangroves the habitat fragmentation has contributed to area reduction and increased edge extension, as well as other anthropogenic consequences that can change abiotic conditions, interfere with ecosystem functioning and lead to the loss of biodiversity. These impacts may affect terrestrial and marine invertebrates living in this coastal system. This study investigated the effects of fragment characteristics (size and matrix type) and distance to edge on ant richness, occurrence of functional groups and of the most frequent species in mangroves in North-Eastern Brazil. Our research covered ten mangrove fragments. We used sardine and honey baits in twelve randomly selected points per area. Twenty-five species of ants were recorded, with Camponotus arboreus, Crematogaster erecta, Azteca sp. and Neoponera villosa being the most frequent. Only four functional groups were found, the most representative of which were the Arboreal Omnivores with Massive Recruitment and the Generalist Predators. No relationship was found between the response variables and the environmental characteristics. Mangroves ants are mostly arboreal; thus, the pattern of competition between ant species or even tree architecture elements may have stronger effects than those related to anthropization, loss of area and fragmentation in this environment. Furthermore, the harsh physical conditions in the mangrove serve as barriers for most exotic species and to some functional groups of ants. This study highlights the natural restrictive nesting and foraging conditions in mangrove areas as the powerful forces in ant community structuring, possibly even more than anthropization itself.     

Direct versus indirect effects of habitat fragmentation on community patterns in experimental landscapes

Habitat area and fragmentation are confounded in many ecological studies investigating fragmentation effects. We thus devised an innovative experiment founded on fractal neutral landscape models to disentangle the relative effects of habitat area and fragmentation on arthropod community patterns in red clover (Trifolium pratense). The conventional approach in experimental fragmentation studies is to adjust patch size and isolation to create different landscape patterns. We instead use fractal distributions to adjust the overall amount and fragmentation of habitat independently at the scale of the entire landscape, producing different patch properties. Although habitat area ultimately had a greater effect on arthropod abundance and diversity in this system, we found that fragmentation had a significant effect in clover landscapes with ≤40 % habitat. Landscapes at these lower habitat levels were dominated by edge cells, which had fewer arthropods and lower richness than interior cells. Fragmentation per se did not have a direct effect on local-scale diversity, however, as demonstrated by the lack of a broader landscape effect (in terms of total habitat area and fragmentation) on arthropods within habitat cells. Fragmentation—through the creation of edge habitat—thus had a strong indirect effect on morphospecies richness and abundance at the local scale. Although it has been suggested that fragmentation should be important at low habitat levels (≤20–30 %), we show that fragmentation per se is significant only at intermediate (40 %) levels of habitat, where edge effects were neither too great (as at lower levels of habitat) nor too weak (as at higher levels of habitat).     

Negative effects of habitat degradation and fragmentation on the declining grassland plant Trifolium montanum

Changes in land use have resulted in a strong decline in the plant diversity of nutrient-poor grasslands, but little is known about the combined effects of habitat degradation and fragmentation on populations of individual species. We studied these effects on stage structure, recruitment, reproduction and offspring fitness in populations of the declining perennial grassland plant Trifolium montanum in central Germany. Density and survival probability of juvenile plants decreased with light competition, measured as leaf area index (LAI) above T. montanum plants, resulting in aged populations with few juvenile plants at unmanaged sites with higher LAI. Reproduction of T. montanum was not related to LAI, but increased strongly with local density, suggesting pollinator limitation in fragmented populations with a low density of flowering plants. In the common garden, the survival of sown offspring increased with mean seed size, whereas seed production of offspring decreased with isolation, and in strong contrast to previous studies, also decreased with size and density of the population of origin. This could be due to increased inbreeding because of pollination between closely related neighbouring plants in dense and large populations. Our results indicate that both habitat degradation and fragmentation have negative effects on populations of T. montanum, but affect different phases of the life cycle. In the short term, the effects of habitat degradation are more important than those of fragmentation, and populations of T. montanum are primarily threatened by an increase in light competition in unmanaged sites, which rapidly affects the dynamics of the populations. The observed opposite effects of habitat fragmentation on reproduction and offspring fitness indicate that the effects of population size, density and isolation on plant fitness and population viability may be complex.     

Conservation of remnant populations of Colchicum autumnale – The relative importance of local habitat quality and habitat fragmentation

Semi-natural habitat is extremely vulnerable to habitat fragmentation and degradation since its socio-economic value has decreased substantially during the last century in most parts of Europe. We evaluated the relative effects of habitat fragmentation and local environmental conditions on population structure and reproductive performance of the long-lived corm geophyte Colchicum autumnale in 17 highly fragmented populations. Habitat isolation did not affect patch occupancy, population structure or plant performance. In contrast, population size and local environment strongly affected population structure and reproductive performance. Densities of all life stages increased with increasing population size. Large populations also showed a higher reproductive performance and a larger proportion of new recruits. Relationships with local growth conditions pointed towards the importance of an open grassland sward for flower and fruit set and the presence of microsites for successful sexual recruitment. These results suggest that the distribution of C. autumnale consists of an assemblage of basically unconnected populations that are remnants of formerly larger populations. This is in accordance with the species' ability to grow clonally, allowing long-term persistence under deteriorating conditions that occurred during a long period of habitat fragmentation. In conclusion, our results indicate that local habitat and population size are more important than habitat fragmentation (i.e. calcareous grassland isolation and surface area) and argue in favour of a management that is primarily focused on local habitat restoration. This is preferentially accomplished by reintroducing grazing practices, complemented by regular setback of spontaneous succession towards forest.