Woodland birds in patchy landscapes: the evidence base for strategic networks

Habitat creation and management within wooded networks is a potentially effective strategy to reduce ecological isolation and the deleterious effects of fragmentation. However, questions remain over the relative advantages of different approaches, e.g. buffering patches vs. increasing connectivity. Potential effects of woodland fragmentation include reduction in regional woodland cover, reduced patch size, edge effects with loss of core habitat, and increased isolation with disruption of dispersal and metapopulation dynamics. We adopt an evidence-based approach to review how each of these affects woodland birds with an emphasis on studies from the UK and use this to identify management priorities for mitigation. There is evidence for both patch area and composition effects: larger woodlands support more woodland bird species, and woods located within sparsely wooded landscapes are less valuable to specialist woodland species. Bird assemblages show a nested pattern with respect to area, and thus species found in small woods also occur in large woods but not vice versa. However, small woods may be preferred by a few edge species, while small woods also have greater variability in bird species composition. Consideration of the metapopulation dynamics of specialist species with poor dispersal shows that creating or buffering large woodlands is more efficient than a greater total area of small fragments. Connectivity appears most useful for widespread generalist species with almost continuous populations. Woodland structure and quality are of overwhelming importance: as well as mature woodland, young growth, scrub and edges are also key components. There is an urgent need to examine the relationship between nest predation and landscape structure within UK woodlands.     

Wooded habitat edges as refugia from microtine herbivory in tallgrass prairies

Meadow voles, Microtus pennsylvanicus , affect the species composition, distribution, and succession of plants in grassland ecosystems, but the effects of voles on herbaceous plants when grasslands are bordered by wooded edges is not known. We investigated the impact of wooded edges on vole distribution and herbivory of relatively palatable and unpalatable native prairie plant species by studying five reconstructed tallgrass prairies with wooded edges in central Iowa. A 50×50 m trapping grid at each site was established to determine the proportion of voles captured at various distances from the edge. We found that meadow voles were less abundant at wooded edges and, in general, increased in number toward the prairie interior. Seedlings of purple prairie clover ( Dalea purpurea ) and Illinois bundleflower ( Desmanthus illinoensis ), a relatively palatable and unpalatable species, respectively, were transplanted onto simulated gopher mounds 2, 5, 10, 20 and 30 m from the edge. The number of plants grazed per species per mound was determined 1 week and 4 weeks after planting. The amount of herbivory on both species was significantly different by distance, with fewer plants eaten 2 m from the edge. Interestingly, the amount of herbivory on relatively unpalatable plants did not differ from more palatable plants. Herbivory on both plant species also varied by site, such that sites with lower vole density tended to have lower amounts of herbivory. These results indicate that wooded edges do have an effect on meadow vole distribution and native prairie seedling herbivory. Because voles avoid wooded edges, seedlings of any species may experience a small refuge from herbivory along wooded edges.     

Influence of edge effects on common vole population abundance in an agricultural landscape of eastern France

Vole demographics are often modified close to wooded environments. Population monitoring in such settings reveals substantial edge effects, although these have seldom been quantified. This study compares the abundance of common voleMicrotus arvalis (Pallas, 1778) populations at various distances from three types of wooded environment (hedgerow, copse and forest) and in two types of habitat (temporary and permanent grassland). In temporary grassland, substantial edge effects were detected along the boundaries of forests and hedgerows. At 100 m from the forest edge, voles were twice more abundant than at 50 m, four times more than at 25 m and eight times more than at 5 m. In permanent grassland, the edge effect was such that voles were practically absent up to 100 m from the forest edge. For both types of grassland, the edge effect was more pronounced for forest habitat than for hedgerows, that is, for elements with a large surface area of wooded environment. Our results suggest that landscape context may greatly affect the local abundance of voles. Given the considerable damage done byM. arvalis, the presence of wooded patches in grassland areas leads to lower vole densities in adjacent areas and this might be an alternative method to controlM. arvalis as a pest species.     

Environmental change and disease dynamics: effects of intensive forest management on Puumala hantavirus infection in boreal bank vole populations

Intensive management of Fennoscandian forests has led to a mosaic of woodlands in different stages of maturity. The main rodent host of the zoonotic Puumala hantavirus (PUUV) is the bank vole (Myodes glareolus), a species that can be found in all woodlands and especially mature forests. We investigated the influence of forest age structure on PUUV infection dynamics in bank voles. Over four years, we trapped small mammals twice a year in a forest network of different succession stages in Northern Finland. Our study sites represented four forest age classes from young (4 to 30 years) to mature (over 100 years) forests. We show that PUUV-infected bank voles occurred commonly in all forest age classes, but peaked in mature forests. The probability of an individual bank vole to be PUUV infected was positively related to concurrent host population density. However, when population density was controlled for, a relatively higher infection rate was observed in voles trapped in younger forests. Furthermore, we found evidence of a "dilution effect" in that the infection probability was negatively associated with the simultaneous density of other small mammals during the breeding season. Our results suggest that younger forests created by intensive management can reduce hantaviral load in the environment, but PUUV is common in woodlands of all ages. As such, the Fennoscandian forest landscape represents a significant reservoir and source of hantaviral infection in humans.     

Dampening of population cycles in voles affects small mammal community structure,decreases diversity,and increases prevalence of a zoonotic disease

Long‐term decline and depression of density in cyclic small rodents is a recent widespread phenomenon. These observed changes at the population level might have cascading effects at the ecosystem level. Here, we assessed relationships between changing boreal landscapes and biodiversity changes of small mammal communities. We also inferred potential effects of observed community changes for increased transmission risk of Puumala virus (PUUV) spread, causing the zoonotic disease nephropatica epidemica in humans. Analyses were based on long‐term (1971–2013) monitoring data of shrews and voles representing 58 time series in northern Sweden. We calculated richness, diversity, and evenness at alpha, beta, and gamma level, partitioned beta diversity into turnover (species replacement) and nestedness (species addition/removal), used similarity percentages (SIMPER) analysis to assess community structure, and calculated the cumulated number of PUUV‐infected bank voles and average PUUV prevalence (percentage of infected bank voles) per vole cycle. Alpha, beta, and gamma richness and diversity of voles, but not shrews, showed long‐term trends that varied spatially. The observed patterns were associated with an increase in community contribution of bank vole (Myodes glareolus), a decrease of gray‐sided vole (M. rufocanus) and field vole (Microtus agrestis) and a hump‐shaped variation in contribution of common shrew (Sorex araneus). Long‐term biodiversity changes were largely related to changes in forest landscape structure. Number of PUUV‐infected bank voles in spring was negatively related to beta and gamma diversity, and positively related to turnover of shrews (replaced by voles) and to community contribution of bank voles. The latter was also positively related to average PUUV prevalence in spring. We showed that long‐term changes in the boreal landscape contributed to explain the decrease in biodiversity and the change in structure of small mammal communities. In addition, our results suggest decrease in small mammal diversity to have knock‐on effects on dynamics of infectious diseases among small mammals with potential implications for disease transmission to humans.     

Habitat preferences of the bank voleMyodes glareolus in a Mediterranean mountain range

The bank voleMyodes glareolus Schreber, 1780 is a widely distributed rodent in Europe, being numerically dominant in small mammal communities living in temperate woodlands. However, it becomes scarce in southern Europe (Mediterranean area) where it reaches the southernmost limit of its distribution range. We studied the habitat preferences of bank voles in 9 plots in a transitional area between Mediterranean and Eurosiberian regions within a Mediterranean mountain. During the study period we captured 1919 small mammals of 9 species, including 287 bank voles (14.95%). Mean density ranged from zero individuals per plot (1.1 ha) at the boreo-subalpine scrubland to 10.27 ± 2.84 (SE) at a Mediterranean river woodland. Statistical path analysis was used to investigate relationships between mean bank vole density and climate and vegetation structure measured within plots. The variables selected by the structural equation model were those related to forest structure, like tree cover and height, dead vegetation, moss, and rock cover. Habitat moisture was also important (microclimatic conditions). Mean climate conditions (and elevation) did not have any significant effect on mean bank vole density, and no significant association with understorey vegetation (eg shrub and herbaceous cover) was observed. Our results pointed out that bank voles were habitat specialists in our study area, being more abundant and frequent in moist woodlands, and rare or absent in shrublands and grasslands.     

Vole diet on experimentally managed reforestation areas in northern Sweden

In northern Sweden two field experiments with the reforestation techniques soil scarification, ploughing, burning and grass herbicidal treatment were performed. Small rodents were trapped regularly on the managed plots and their stomachs were examined microscopically for diet composition. Both bank voles Clethrionomys glareolus and field voles Microtus agrestis were common on the reforestation areas while only a small number of grey-sided voles Clethrionomys rufocanus were taken. All three species underwent a population cycle during the studies. The management techniques generally resulted in small and irregular effects on the food selection. The most pronounced changes were lower intake of grasses by M. agrestis after herbicidal treatment and of filamentous tree lichens by C. glareolus after most treatments.
Both bank voles and field voles ate predominantly forbs in the summer half of the year, whereas the field voles took also a considerable amount of grass. As a complement to green vegetable-matter bank voles ate berries and fungi in summer-autumn and tree lichens at other times of the year, but seeds and animals food only in very small amounts. Ail three species consumed large quantities of dwarf-shrubs in autumn and especially in winter. Considerable amounts of bark were eaten by field voles and a smaller proportion by bank voles in autumn-winter.
Both for bank and field voles there were indications of worsening food conditions as the population cycle went on, There were, for example, an increase in grass and bark intake in field voles and a decrease in seeds and berries for the hank vole.     

Beyond the Mediterranean peninsulas: evidence of central European glacial refugia for a temperate forest mammal species, the bank vole (Clethrionomys glareolus)

This study details the phylogeographic pattern of the bank vole, Clethrionomys glareolus, a European rodent species strongly associated with forest habitat. We used sequences of 1011 base pairs of the mitochondrial DNA cytochrome b gene from 207 bank voles collected in 62 localities spread throughout its distribution area. Our results reveal the presence of three Mediterranean (Spanish, Italian and Balkan) and three continental (western, eastern and 'Ural') phylogroups. The endemic Mediterranean phylogroups did not contribute to the post-glacial recolonization of much of the Palaearctic range of species. Instead, the major part of this region was apparently recolonized by bank voles that survived in glacial refugia in central Europe. Moreover, our phylogeographic analyses also reveal differentiated populations of bank voles in the Ural mountains and elsewhere, which carry the mitochondrial DNA of another related vole species, the ruddy vole (Clethrionomys rutilus). In conclusion, this study demonstrates a complex phylogeographic history for a forest species in Europe which is sufficiently adaptable that, facing climate change, survives in relict southern and northern habitats. The high level of genetic diversity characterizing vole populations from parts of central Europe also highlights the importance of such regions as a source of intraspecific genetic biodiversity.     

Seasonal and annual variations in the prevalence of helminths in a cyclic population of Clethrionomys glareolus

Six cestode and two nematode species were recovered from a cyclic population of Clethrionomys glareolus in southern Norway. The cestode Aprostatandrya macrocephala and the nematode Heligmosomum glareoli were euconstant species which displayed late spring/early summer peaks in numbers after which they declined. This decrease in prevalence was correlated with decreasing mean age of the host population. Increases in the prevalence of infections with the nematode Syphacia nigeriana in bank voles were correlated to increasing numbers of Microius agrestis in the mixed woods. Cestode larvae were at maximum prevalence during peak years of bank vole abundance. Pathological conditions such as ascites and peritoneal adhesions, as well as splenic and adrenal hypertrophy, were associated with cestode larvae. There was no general trend of an increase in the prevalence of helminth infection with increasing population density of the bank voles.     

How important are large vs. small forest remnants for the conservation of the woodland flora in an urban context?

Aim To test whether, in an urban area, small forest fragments are more important than large ones, the present study aimed at firstly exploring the relationship between plant functional groups or individual species and urban woodlot characteristics such as patch area and isolation, and secondly investigating whether equal‐sized combinations of islands tend to differ in species richness. Location The city of Brussels. Methods We considered the relationship between size, species richness and plant functional groups among one very large (1666 ha) and 11 rather small (2–123 ha) woodlots. Results The largest woodlot harboured species missing in the smaller ones. The species‐area relationship plotted for these wooded patches fitted the semilog model very well. Twenty‐three species had a significantly higher frequency in the main forest. Only six species had a significantly higher frequency in smaller woodlots. The occurrence of species groups with high conservation value (e.g. ancient forest species, rare species) was higher in large patches. However, a SLOSS analysis showed that habitat subdivision appeared to be associated with increased species counts. A marginally nonsignificant effect of distance to the main forest became significant when matrix species were removed from the patch samples. Conclusions Although overall the data showed a higher conservation value for large woods, some plant functional groups (e.g. woodland species vs. ancient forest species) responded differently to fragmentation. This illustrates that, for conservation strategies, studies considering the biotic characteristics of remnants should focus on the species number of particular plant functional groups, especially those with high conservation value. Furthermore, matrix species should be removed from the analysis in order not to mask underlying patch size and distance effects.     

Does removal of an alien predator from small islands in the Baltic Sea induce a trophic cascade?

Changes in carnivore abundance can alter the distribution and abundance of plants on a community wide basis, an effect known as a trophic cascade. Because alien predators can have a disproportionate impact, compared to native predators, on herbivore populations, they may induce stronger trophic cascades in plant communities than native predators. We studied the indirect effects of the removal of an alien predator, the American mink Mustela vison on plant communities on small islands in the Baltic Sea, SW Finland. Mink had been removed from a group of islands for 12 yr, while another group of islands with mink presence served as a control area. Field voles Microtus agrestis and bank voles Myodes glareolus exert strong grazing pressure on the island vegetation and are an important part of mink diet. On nine islands of the mink removal area and five islands of the control area we studied the vegetation in ten randomly chosen plots; five in herbaceous and five in woody (i.e. dwarf shrub) vegetation. We studied the cascading impacts of mink predation on grassy and woody vegetation using the Shannon diversity and equitability indices and comparing abundances of different species between mink removal and control islands. Diversity and equitability of plant communities were higher on mink removal islands. In grassy patches, abundances of several species differed between mink removal and control islands. Our results demonstrate, for the first time, that alien predator removal may induce a trophic cascade on small islands.     

Regulation of forest defoliating insects through small mammal predation: reconsidering the mechanisms

Population densities of forest defoliating insects may be regulated by small mammal predation on the pupae. When outbreaks do occur, they often coincide with warm, dry weather and at barren forest sites. A proposed reason for this is that weather and habitat affect small mammal population density (numerical response) and hence pupal predation. We propose an alternative explanation: weather and habitat affect small mammal feeding behaviour (functional response) and hence the outbreak risks of forest pest insects. We report results from laboratory and field-enclosure experiments estimating rates of pupal predation by bank voles (Myodes glareolus) on an outbreak insect, the European pine sawfly (Neodiprion sertifer), at different temperatures (15 and 20 °C), in different microhabitats (sheltered and non-sheltered), and with or without access to alternative food (sunflower seeds). We found that the probability of a single pupa being eaten at 20 °C was lower than at 15 °C (0.49 and 0.72, respectively). Pupal predation was higher in the sheltered microhabitat than in the open one, and the behaviour of the voles differed between microhabitats. More pupae were eaten in situ in the sheltered microhabitat whereas in the open area more pupae were removed and eaten elsewhere. Access to alternative food did not affect pupal predation. The results suggest that predation rates on pine sawfly pupae by voles are influenced by temperature- and habitat-induced variation in the physiology and behaviour of the predator, and not necessarily solely through effects on predator densities as previously proposed.     

Small habitat islands are inferior breeding habitats but are used by some great tits – competition or ignorance?

Great tit breeding performance in small habitat islands in an agricultural landscape was compared to that in large deciduous woods. Clutch size was similar in both habitats, but more nestlings starved and fledgling weight was lower in the small habitat islands. The area of wooded habitat in the territories of tits breeding in the small habitat islands was less than what is common to encounter in optimum habitat, deciduous woods. I suggest that this may be responsible, ultimately or proximately, for the poor performance in the small habitat islands. Two possible mechanisms for this effect are discussed. It is possible that the small habitat islands were perceived as inferior by the tits and used by competitively inferior individuals that also were less successful in raising a brood. However, the results were obtained even when female age and weight were controlled for. Therefore, the competitive ability (and probably also young raising competence) may be the same for tits breeding in small habitat islands as for those breeding in optimum habitat. Such a pattern is conceivable if tits have been selected for maintaining a large territory as a response to the close presence of other tits and not as a means of securing a large feeding area. Historically, territories large enough to reduce predation or mate competition (or any cause of territoriality) may always have contained ample food. The presence of small islands, without close neighbours but with insufficient food, may be evolutionarily new. It is probably typical of modern agricultural landscape.     

Effect of forest fragmentation on dung beetle communities and functional consequences for plant regeneration

I compared dung beetle communities and assessed some of their functional effects (dung removal, seed burial, seedling establishment) in continuous forest with those in 1-ha and 10-ha forest fragments in Central Amazonia. I followed the fate of seeds until seedling establishment for three native tree species, using clean seeds and seeds surrounded by dung. The 1-ha fragments had half the number of dung beetle species captured in continuous forest and in 10-ha fragments. The continuous forest sites and the 1-ha fragments had similar number of individuals, but in the 10-ha fragments dung beetles were twice as abundant. Mean beetle size increased with increasing forest area. Dung removal and seed burial rates were higher in continuous forest than in forest fragments. Seed predation rates were higher in the forest fragments. In all sites, the proportion of seedlings established from seeds surrounded by dung vs clean seeds was the same, and it was the same in continuous forest vs fragments. When comparing seeds that remained on the forest floor with seeds buried by dung beetles, a higher percentage of seedlings established from the latter. Conservation programs that aim to maintain the regeneration ability of forest fragments must incorporate all the important components involved in seedling establishment; in Central Amazonia these include dung beetles as secondary dispersers. It is important that studies start measuring directly not only the first-order effects of forest fragmentation on species, but also the higher-order functional effects.     

Area mediated shifts in bird community composition: a study on a fragmented Mediterranean grassland

The effects of habitat fragmentation on birds have often been studied in forest specialist species. Here we aimed at comparing the response of open habitat birds within a range of habitat specialization. The study area was a Mediterranean pseudo-steppe, designated as important for conservation yet fragmented by tree encroachment. We defined bird species dependency on steppe-like habitat by a correspondence analysis, allowing us to distinguish between specialists, generalists and scrubland species. We studied species abundance in relation to fragment area, testing whether species representation in fragments differed from those in continuous habitat. This analysis showed a contrasted response to fragment size between “open habitat” specialist species and generalist ones. Open habitat species were under-represented in the smallest fragments, while generalist were over-represented in small fragments in comparison to their distribution in continuous habitats. We discuss how these results can be linked to species habitat requirements. We find that scrubland species seem to be favoured by encroachment of woody vegetation, as they are able to explore and use the wooded matrix; however specialist species are restricted to open patches and are sensitive to a reduction in patch size. This allows us to predict how different species can exhibit a different sensitivity to habitat fragmentation.     

Natural infections of small mammals with blood parasites on the borderland of boreal and temperate forest zones

Blood parasites of small mammals living in Białowieża Forest (eastern Poland) were investigated between 1996 and 2002. The following haemoparasite species were found:Trypanosoma (Herpetosoma) evotomys in bank voleClethrionomys glareolus; T. (H.) microti in root voleMicrotus oeconomus; Babesia microti in root vole;Hepatozoon erhardovae in bank vole andHepatozoon sp. in root vole. Some non-identifiedBartonella species were found in bank vole, root vole, field voleMicrotus agrestis, yellow-necked mouseApodemus flavicollis, common shrewSorex araneus, Eurasian water shrewNeomys fodiens, and Mediterranean water shrewN. anomalus. The prevalence and diversity of blood parasites were lower in shrews than small rodents. Totally, 52.0% of bank voles, 50.0% of root voles, 32.5% of common shrews, and 41.2% of Eurasian water shrews were infected with any of the blood parasites. Mixed infections were seldom observed in bank vole (17.3% of investigated individuals) and root vole (14.7%). No animals were infected with three or four parasites simultaneously. Infection of Białowieża small mammals with haemoparasites seemed to be similar to those described in other temperate forest regions rather than boreal ones. Infection rates of rodent species seem to be higher in their typical habitats: for bank vole it was the highest in mixed forest, whereas for root vole in sedge swamp. The results suggest that Arvicolidae play a greater role than Muridae or Soricidae in maintenance ofBabesia andHepatozoon foci in natural environments of central Europe.     

Spatio-temporal patterns of habitat use in voles and shrews modified by density, season and predators

1.?Although the intrinsic habitat preferences of a species can be considered to be fixed, the realized habitat use depends on the prevailing abiotic and biotic conditions. Often the core habitats are occupied by dense and stable populations, while marginal habitats become occupied only at times of high density. In a community of interacting species, habitat uses of different species become inter-related, for example an increased density of a strong competitor forcing a weaker competitor to use more marginal habitats. 2.?We studied the spatio-temporal distribution patterns of three common small mammal species, the bank vole Myodes glareolus; the field vole Microtus agrestis; and the common shrew Sorex araneus, in a 4-year trapping study carried out on six large islands, each containing a mixture of three main habitat types (forest, field and clear-cut). We experimentally released least weasels (Mustela n. nivalis) to some of the islands to see how the focal species respond to increased predation pressure. 3.?Both vole species were largely restricted to their core habitats (bank voles to forests and field voles to fields) at times of low population density. With increasing density, the relative habitat use of both species increased in the clear-cut areas. The common shrew was a generalist in its habitat use at all population densities. 4.?The release of the weasels changed the habitat use of all study species. 5.?The vole species showed a stronger aggregated pattern than the common shrew, especially at low population density. The vole aggregations remained in the same localities between seasons, except in the case of bank voles after the weasels were released. 6.?Bank voles and field voles avoided each other at high density. 7.?We conclude that intrinsically differential habitat requirements and flexibility to modify habitat use facilitate the coexistence of the two competing vole species in mosaic landscapes consisting of boreal forests and open habitats.     

Structure and seasonal dynamics of the biotic community ectoparasites of the bank vole in the Il'men'-Volkhov lowland

A biodiversity, seasonal dynamics and parasite load of a single individual and local population of the bank vole (Clethrionomys glareolus) were studied in coniferous and mixed forests of the Ilmen-Volkhov lowland (neighborhood of Oskuy village, Chudovo region, Novgorod Province) in the period from June 1999 till May 2002. The Gero's traps were used for collecting the host. Lines of traps stood in each place during 3-5 days and were checked twice a day. Trapping of micromammalian hosts and collecting of parasites took place each month, except a few gaps. Total number of collected animals 2854 including 1405 bank voles. The 29 ectoparasite species were recorded on the bank vole in the area of study. Among some mites and fleas a few species are accidental parasites probably accepted by voles from other species of animals. In the Oskuy area, the bank voles are the main and additional hosts of 25 ectoparasite species: fleas--8, lice--1, ixodid ticks--2, gamasid mites--7, acariform mites 7. Species composition of ectoparasites, their occurrence and abundance change during the year. Seasonal changes of abundance and occurrence indices are most expressed in the temporary ectoparasites (ixodid ticks, chiggers, gamasid mites), while in the permanent parasites (lice, acariform mites: Myobiidae, Myocoptidae, Listrophoridae), the seasonal fluctuations of indices are displayed in a less scale. Most vole specimens were infected with this or that ectoparasite species. The parasitocenosis on an individual specimen usually included less than 10 species of the total number 29 species recorded on the bank vole in the area investigated. One ectoparasite species was recorded on 21% of host specimens, 2-5 ectoparasite species were found on 71% of host individuals. Maximal number of ectoparasite species (10 species) was registered on one specimen only. The parasite load was dispersed unevenly among the infected voles. Mean number of parasites of all species on a host individual varied from 124 to 295. The highest grade of parasites (237-297 parasite specimens) was found in the voles with 7-10 species of parasites.     

Changes in species composition following field and tree layer restoration and management in a wooded hay meadow

A hillside with abandoned, wooded hay meadows in Leikanger in Sogn og Fjordane county, Western Norway, was investigated in 1992–1995. A management regime combining restoration (topping) of old pollarded trees, scything and removal of hay was reintroduced in the area. The availability of light, hydrological conditions and the nutrient content of the soil were the most important factors determining development of hay meadow vegetation to forest vegetation. Indicators of shaded conditions included Valeriana sambucifolia and Oxalis acetosella , while Anthriscus sylvestris was indicator of light conditions. Species such as Campanula rotundifolia, Pimpinella saxifraga, Plantago lanceolata, Lychnis viscaria and Hieracium pilosella , which were recorded mostly on dry ridges, were regarded as "relict" species, more common formerly when the area was managed in a traditional way. The cover of tall grasses and herbs, such as Dactylis glomerata, Poa trivialis and Anthriscus sylvestris , increased the first two years after restoration. This indicates an increase in available nutrients, probably a "pioneer clearing fertilization" due to the introduced pollarding. Restoration and management resulted in drier, more homogenous ecological conditions and reduced the differences in local climate between formerly open meadows and more wooded areas in the restored area. However, most of the vegetation seemed to be quite stable during the study period. One class, however, comprised releves from several unstable plots of the experimental site. The frequency of some light-demanding species increased in the experimental site, while the frequency of Oxalis acetosella and Epilobium montanum increased in the control site.