Patterns of Clonal Growth Modes Along a Chronosequence of Post-Coppice Forest Regeneration in Beech Forests of Central Italy

Forest coppicing leads to changes in composition of the herbaceous understory through soil disturbance and alteration of the light regime. While the role of seed dispersal traits at the start of succession after coppicing has been extensively studied, the role of persistence traits such as clonal growth and bud banks is not yet sufficiently understood. To gain better understanding of this role, we studied the patterns of clonal growth organs and related clonal traits of species in a series of coppiced beech forests of the Central Apennines (Marches region, Italy) in various stages of recovery after the last coppicing event. We conducted stratified random sampling and established a chronosequence of recovery stages based on stand age (reflecting the number of years since the last coppicing). The beech stands were classified into three age groups (Post-logged, Recovering, and Old-coppice stands) according to the characteristic stages of beech coppice dynamics. Clonal growth organs and the corresponding clonal traits of plants in the forest understory vegetation were assessed with the help of a CLO-PLA1 database. We found no significant change in the proportion of clonal species along the studied chronosequence. In contrast, most of the traits and about the half of the clonal growth organs showed correlation with stand age or preference for a certain habitat (i.e., stage of regeneration). Clonal and bud bank traits proved to play an important role in the persistence of species subjected to forest coppicing cycles in the studied area.     

Biodiversity implications of coppice decline,transformations to high forest and coppice restoration in British woodland

Coppice systems are amongst the earliest forms of woodland management known, and on some sites, their use has been documented for centuries. Distinctive assemblages of plants and animals are associated with such systems and are highly valued in nature conservation terms. The richness of such assemblages, and conversely, the species that do not thrive under coppice, are linked to the alternation of relatively short light and dark phases, and the juxtaposition of stands at different stages in the coppice cycle. Vascular plants in the ground flora, invertebrates of open glades and scrub, and small birds of the understorey may have become more abundant in coppice than they would have been under ‘natural’ forest conditions. By contrast, epiphytes dependent on mature trees and species of large-sized deadwood are less favoured by coppice management. Coppice systems developed to meet the local community needs. As social and economic conditions changed, so coppicing declined and the woods were transformed into high forest through neglect or deliberate management. High forests differ from coppice stands in their spatial and temporal dynamics, and consequently, in their wildlife, particularly with respect to their vertical structure pattern, extent of open space and young growth, spatial heterogeneity, tree and shrub composition, and browsing levels. Three issues for the conservation of biodiversity arise from these changes: (1) What priority and resources should be given to halting further decline, by maintaining coppice compared to allowing sites to develop with more ‘natural’ high forest structures and dynamics; will associated high-forest species recolonize? (2) If we restore coppice systems, will the species assemblages present in the past also recover, under current and future changes in environmental conditions; i.e is the transformation reversible under current environmental conditions? (3) Are there other ways in which ‘coppice-associated’ species might be maintained? We identify research gaps and proposals to address these issues.     

Dynamics of herbaceous vegetation during four years of experimental coppice introduction

Understanding the effects of coppicing on forest ecosystems is important for progress towards sustainable forest management. A newly established coppicing experiment in a secondary temperate deciduous forest in the SE Czech Republic provides a rather unique insight into succession driven by canopy thinning in a forest still lacking species typical for forests established a long time ago. Herbaceous layer vegetation was monitored for four subsequent years in 2012–2015. We focused on the influence of canopy thinning intensity in two different forest types defined by dominant tree species (oak and lime). Our results showed that the opening of the canopy had immediate effects on herbaceous vegetation. Coverage, species richness and compositional patterns followed the coppicing intensity gradient. The dominant tree species had contrasting effects. Under oak, the reaction to coppicing was weak. Under lime, strong reaction both related to coppicing intensity and temporal development was observed. Herbs with short life cycle had the greatest contribution, but perennial grasses also began to increase their coverage after coppicing. Several invasive species, mostly short-lived herbs, emerged but are supposed to retreat as the succession will proceed. We conclude that coppice introduction into a secondary forest led to contrasting patterns related to dominant tree species. The marked difference was probably due to the slow succession towards a future forest community saturated by species. This process may now be further diversified by coppicing management.     

Plant species diversity in Mediterranean old-growth forests: A case study from central Italy

Abstract

To investigate the differences in understorey composition and diversity between old-growth and managed forests, we analyzed an old-growth and a managed beech stand in the same area displaying similar abiotic features. We considered variations in understorey species composition and richness. The sampled understorey species were characterized in terms of functional traits, Ellenberg's indicator values and taxonomic distinctness; next, we calculated four different pairwise plot-to-plot dissimilarity matrices based on species composition, functional traits, Ellenberg's indices and taxonomic distances. We applied a permutational multivariate extension of ANOVA to test whether the forest stands significantly differ in the considered features. Indicator values of all plant species in managed and old-growth stands were evaluated.

The old-growth forest had a higher species richness; permutational analysis of variance showed significant differences between the two stands in plant species composition, functional traits, Ellenberg indices and taxonomic distances. Indicator species analysis highlighted 14 indicator species for the unmanaged stand, while only 3 indicators were found for the managed one.

The results suggest that forest management determines ecological differences that strongly affect plant species composition.

The knowledge of natural stands dynamics could allow development of new approaches and practices in forest management focusing on biodiversity conservation.     

Changes of vascular plant diversity along a chronosequence of beech coppice stands,central Apennines,Italy

Abstract

This research sought to understand the patterns of vegetation recovery after disturbances because of coppice management in beech forests. Eighty sampling units from the mountain belt of the Marche region (Apennines, Italy) were collected according to a stratified sampling based on their geological setting (limestone, sandstone), elevation classes and age after last coppicing (to represent a chronosequence, from 1 to 90 years). The expected successional trend of decreasing species richness was confirmed, together with some stabilisation processes for older stand ages. However, more complex diversity patterns were found when total species richness was decomposed into the species richness of five social behaviour types (SBTs), defined based on the species' habitat preference. On both bedrocks, temporal gradients explained the observed diversity changes at the stand level: forest specialists increased whereas non-forest species decreased. A relatively long time after coppicing (40–60 years), the contribution of the beech specialist species doubled, whereas non-forest and species from anthropogenic habitats decreased by about 50%. On sandstone, the contribution of gap species also decreased over a long-period, and the beech stands experienced stronger changes over time. We conclude that the decomposition of total species richness in terms of SBT affords the opportunity to identify temporal references for thresholds which can be used to assess plant diversity status in relation to management schedules and conservation policy decisions.     

Patterns of seedling recruitment in West-Mediterranean Quercus ilex forest influenced by canopy development

Abstract. Quercus ilex (holm oak) coppice forests belong to the most representative communities in the western part of the Iberian Peninsula. Due to the high sprouting potential of holm oak after coppicing, the dynamics of recruitment of new individuals in existing populations has been largely overlooked. In these forests, the density of seedlings increases in old stands, as a result of the remarkable increase in the recruitment of 1-yr and 2-yr old seedlings, probably due to better environmental conditions for germination. Older seedlings show a tendency towards increasing survival under some degree of canopy closure, since their density appears to be higher just before complete canopy closure takes place. Nevertheless, the reduced growth of seedlings in older stands due to low light levels causes age distributions of holm oak seedlings to change drastically, with saplings older than 15 yr becoming very unusual in these stands. Therefore, a bottleneck of recruitment appears at this sapling stage and prevents vigorous regeneration just before thinning. Present conditions of genet density and management tend to prevent population renewal. Dynamics of the seedling bank reinforce the role of gap formation through small or large scale perturbations in population turnover.     

Distinguishing between turnover and nestedness in the quantification of biotic homogenization

Compositional changes through local extinction and colonization are inherent to natural communities, but human activities are increasingly influencing the rate and nature of the species being lost and gained. Biotic homogenization refers to the process by which the compositional similarity of communities increases over time through a non-random reshuffling of species. Despite the extensive conceptual development of the homogenization framework, approaches to quantify patterns of homogenization are scarcely developed. Most studies have used classical dissimilarity indices that actually quantify two components of compositional variation: turnover and nestedness. Here we demonstrate that a method that partitions those two components reveals patterns of homogenization that are otherwise obscured using traditional techniques. The forest understorey vegetation of an unmanaged reserve was recorded in permanent plots in 1979 and 2009. In only thirty years, the local species richness significantly decreased and the variation in the species composition from site to site shifted towards a structure with reduced true species turnover and increased dissimilarity due to nestedness. A classic analysis masked those patterns. In summary, we illustrated the need to move beyond the simple quantification of homogenization using classical indices and advocate integration of the multitude of ways to quantify community similarity into the homogenization framework.     

Relationships between the age of northern Kantou plain (central Japan) coppice woods used for production of Japanese forest mushroom logs and butterfly assemblage structure

To discern mechanisms maintaining the diversity of grassland and forest butterflies in coppice woods managed for the production of Japanese forest mushroom logs, we investigated the butterfly fauna in cut-over land tracts shortly after felling and 5 year later, and in forest stands 10, 15, and 25 year after felling (here, we use the term “forests” when referring to the chronosequence of these treed stands). Butterfly species richness and diversity (H′) and the densities of individuals were highest in cut-over lands 5 year after clear-cutting, followed by 25-year-old forest stands. In forests, the richness and densities of forest butterfly species were higher than were those of grassland species. Among forest stands of different ages, forest butterfly species’ richness and the densities of individuals were highest in 25-year-old woods nearing felling time. Some forest butterfly species were observed only in forests. The species richness and densities of grassland butterflies were much higher in cut-over lands 0 and 5 year post felling than in forests; grassland species were rarely found in stands ≥10 year old. Thus, cut-over lands seem to function as temporary habitats for grassland species. Furthermore, the number of forest butterfly species was the same in cut-over lands 5 year after felling and in 25-year-old forest stands; the densities of forest butterfly species was higher in these cut-over lands than in the forest stands. Forest butterfly species living on cut-over land 5 year post felling sipped flower nectar, laid eggs on host plants, and practiced territorial behaviour involved in mate finding. Hence, these cut-over lands functioned as important habitats for various developmental stages of forest butterflies. In conclusion, traditional coppicing in woods for production of Japanese forest mushroom logs is very important for the maintenance of diversity in grassland and forest butterfly species.     

Genetic resilience in a historically profited root sprouting oak (Quercus pyrenaica Willd.) at its southern boundary

Recent land use changes entailed the abandonment of traditional forest practices, which genetic and ecological sustainability should be evaluated in the frame of current socioeconomic and ecological changes. The present study aimed to assess the conservation value of Quercus pyrenaica Willd. in stands subjected to two traditional rural practices, one specific (coppicing) and the other generic (maintenance of open parklands), and their effects on genetic diversity and clonal structure in this singular root sprouting oak at the southern limit of its distribution. Genetic diversity measures of seven nuclear simple sequence repeat markers were compared, and Hardy–Weinberg disequilibria were tested to be originated from recent population demography. Results showed that, regardless of forest structure, the degree of clonality was very similar (～60 %), being allele and lineage density proportional to stem density. Nevertheless, evenness of clonal distribution was higher in coppice, suggesting more homogeneous management than in open woodland. Contrary to previous beliefs, coppice stands do not involve genetic diversity losses; rather, the process of forest conversion into open woodland leads to the removal of numerous genetic lineages and low frequency alleles. The ancient presence of Q. pyrenaica in the region, which constituted quaternary glacial refuge, may contribute to its high genetic diversity. Historical vicissitudes in this anthropogenic deforested territory remark its resilient character; based on a specific fire pre-adaptive trait, continued coppicing fostered the preservation of its natural genetic diversity. This study evidences the importance of the integration of molecular and historical approximations to assess the genetic and conservation status of a secularly profited woody species.     

Removal of cattle accelerates tropical dry forest succession in Northwestern Mexico

Domestic livestock influence patterns of secondary succession across forest ecosystems. However, the effects of cattle on the regeneration of tropical dry forests (TDF) in Mexico are poorly understood, largely because it is difficult to locate forests that are not grazed by cattle or other livestock. We describe changes in forest composition and structure along a successional chronosequence of TDF stands with and without cattle (chronic grazing or exclusion from grazing for ~ 8 year). Forest stands were grouped into five successional stages, ranging from recently abandoned to mature forest, for a total of 2.7 ha of the sampled area. The absence of cattle increased woody plant (tree and shrub) density and species richness, particularly in mid-successional and mature forest stands. Species diversity and evenness were generally greater in sites where cattle were removed and cattle grazing in early successional stands reduced establishment and/or recruitment of new individuals and species. Removal of cattle from forest stands undergoing succession appears to facilitate a progressive and non-linear change of forest structure and compositional attributes associated with rapid recovery, while cattle browsing acts as a chronic disturbance factor that compromises the resilience and structural and functional integrity of the TDF in northwestern Mexico. These results are important for the conservation, management, and restoration of Neotropical dry forests.     

Comparison of vascular plant diversity and species composition of coppice and high beech forest in the Banat region,Romania

Abandonment of coppicing is one of the main reasons of diversity decline in European temperate forests. To reverse this trend, coppicing has been reintroduced in several forests, especially in areas of high conservation value. However, empirical information on the effects of coppicing on plant diversity and composition is still scarce. By comparing vegetation data from active coppices and beech-dominated high forests in the Banat region in Romania, we tested the hypothesis that coppices have a higher plant diversity and different plant species composition than high forests. Data were collected in 60 randomly placed phytosociological relevés and were analysed using linear models and multivariate methods. As expected, we found differences in understorey plant composition. Herb species with various environmental demands–sciophytes-(hemi)-heliophytes and grassland species–were more frequent in coppices whereas high forests had more vernal species and sciophytes. Coppices also had slightly greater plant diversity but did not differ in herb species richness. Our results demonstrate that coppicing has a small positive effect on plant diversity but a large effect on species composition. Coppice restoration may be especially beneficial for populations of thermophilous and non-forest species.     

Carbon accrual rates,vegetation and nutrient dynamics in a regularly burned coppice woodland in Germany

Historically, large areas of forest in Europe were managed as coppice woodland to produce wood‐based fuel for the smelting industry. We hypothesized that this practice produced a legacy effect on current forest ecosystem properties. Specifically, we hypothesized that the historical form of coppicing may have produced a legacy of elevated stocks of soil organic carbon (SOC), nutrients and black carbon (BC) in soil as fire was routinely used in coppiced woodland to clear land. We further hypothesized that these changes in soil properties would result in increased biodiversity. To test these hypotheses, we sampled the surface soil (0–5, 5–10 and 10–20 cm) from a chronosequence of forest sites found in the Siegerland (Germany) that had been coppiced and burned 1, 2, 3.5, 6, 8, 11 and 17 years before present. Mature beech and spruce forests (i.e., >60 years) were also sampled as reference sites: to provide a hint of what might occur in the absence of human intervention. We measured stocks of SOC, BC, NO₃‐N, P, K, Mg, as well as cation exchange and water‐holding capacity, and we mapped plant composition to calculate species richness and evenness. The results showed that coppicing in combination with burning soil and litter improved soil nutrient availability, enhanced biodiversity and increased SOC stocks. The SOC stocks and biodiversity were increased by a factor of three relative to those in the mature beech and spruce forests. This study shows that traditional coppicing practice may facilitate net C accrual rates of 20 t ha^?1 yr^?1 and maintain high biodiversity, indicating that aspects of traditional practice could be applied in current forest management to foster biodiversity and to mitigate climate change.     

Bird Community Changes in Response to Single and Repeated Fires in a Lowland Tropical Rainforest of Eastern Borneo

Our current understanding of bird community responses to tropical forest fires is limited and strongly geographically biased towards South America. Here we used the circular plot method to carry out complete bird inventories in undisturbed, once burned (1998) and twice burned forests (1983 and 1998) in East Kalimantan (Indonesia). Additionally, environmental variables were measured within a 25 m radius of each plot. Three years after fire the number of birds and bird species were similar for undisturbed and burned forests, but species diversity and turnover were significantly lower in the burned forests. The bird species composition also differed significantly between undisturbed and burned forests, with a strong decline of closed forest preferring bird species accompanied by a strong increase in degraded forest preferring species in burned forests. These differences were strongly related to differences in environmental conditions such as shifts in vegetation cover and layering and differences in ground and understorey vegetation structure. We also found significant shifts in body mass distribution, foraging height and feeding guilds between the bird communities in unburned and burned forests. Surprisingly, repeated burning did not lead to increasing impoverishment of the avifauna, and both once and twice burned forests still contained most of the bird species that were also present in undisturbed forest, even though their densities were considerably lowered.     

Forest complexity drives dung beetle assemblages along an edge-interior gradient in the southwest Amazon rainforest

1. The habitat heterogeneity hypothesis predicts that heterogeneous habitats may provide more niches and diverse ways of exploiting environmental resources, thereby allowing more species to coexist, persist and diversify. 2. We aimed to investigate how an edge-interior gradient related to forest complexity influences species composition, abundance and richness of dung beetles in the western Amazon rainforest. We expected dung beetle abundance and richness to increase along the forest edge-interior gradient, in accordance with the habitat heterogeneity hypothesis. We also expected strong changes in species composition driven by species turnover in the forest interior and nestedness along the forest edges. We sampled dung beetles using baited pitfall traps across an edge-interior gradient. We also assessed the variation in forest features along the edge-interior gradient to identify changes in forest complexity. 3. Both species richness and abundance of dung beetles increased along the forest edge-interior, following the gradient of forest complexity. The Sorensen dissimilarity of dung beetle assemblages was higher among sampling units placed near the forest edge, although neither turnover, nor nestedness was different between the extremes of the forest edge-interior gradient. There was a clear compositional change along the edge-interior gradient mostly driven by species turnover. Individual indicator value analysis revealed that species were strongly associated with the forest interior conditions. 4. The simplification of the Amazon rainforest near clearings causes compositional changes in dung beetle assemblages. These changes are characterised by species-poor and low-abundance assemblages and may impair dung beetle ecological functions and therefore forest recovery.     

Effects of Previous Land-Use on Plant Species Composition and Diversity in Mediterranean Forests

At some point in their history, most forests in the Mediterranean Basin have been subjected to intensive management or converted to agriculture land. Knowing how forest plant communities recovered after the abandonment of forest-management or agricultural practices (including livestock grazing) provides a basis for investigating how previous land management have affected plant species diversity and composition in forest ecosystems. Our study investigated the consequences of historical “land management” practices on present-day Mediterranean forests by comparing species assemblages and the diversity of (i) all plant species and (ii) each ecological group defined by species’ habitat preferences and successional status (i.e., early-, mid-, and late-successional species). We compared forest stands that differed both in land-use history and in successional stage. In addition, we evaluated the value of those stands for biodiversity conservation. The study revealed significant compositional differentiation among stands that was due to among-stand variations in the diversity (namely, species richness and evenness) of early-, intermediate-, and late-successional species. Historical land management has led to an increase in compositional divergences among forest stands and the loss of late-successional forest species.     

Food-Web Structure in Relation to Environmental Gradients and Predator-Prey Ratios in Tank-Bromeliad Ecosystems

Little is known of how linkage patterns between species change along environmental gradients. The small, spatially discrete food webs inhabiting tank-bromeliads provide an excellent opportunity to analyse patterns of community diversity and food-web topology (connectance, linkage density, nestedness) in relation to key environmental variables (habitat size, detrital resource, incident radiation) and predators:prey ratios. We sampled 365 bromeliads in a wide range of understorey environments in French Guiana and used gut contents of invertebrates to draw the corresponding 365 connectance webs. At the bromeliad scale, habitat size (water volume) determined the number of species that constitute food-web nodes, the proportion of predators, and food-web topology. The number of species as well as the proportion of predators within bromeliads declined from open to forested habitats, where the volume of water collected by bromeliads was generally lower because of rainfall interception by the canopy. A core group of microorganisms and generalist detritivores remained relatively constant across environments. This suggests that (i) a highly-connected core ensures food-web stability and key ecosystem functions across environments, and (ii) larger deviations in food-web structures can be expected following disturbance if detritivores share traits that determine responses to environmental changes. While linkage density and nestedness were lower in bromeliads in the forest than in open areas, experiments are needed to confirm a trend for lower food-web stability in the understorey of primary forests.     

Understorey plant community structure in lower montane and subalpine forests, Grand Canyon National Park, USA

Aim Our objectives were to compare understorey plant community structure among forest types, and to test hypotheses relating understorey community structure within lower montane and subalpine forests to fire history, forest structure, fuel loads and topography. Location Forests on the North Rim of Grand Canyon National Park, Arizona, USA. Methods We measured understorey (< 1.4 m) plant community structure in 0.1‐ha plots. We examined differences in univariate response variables among forest types, used permutational manova to assess compositional differences between forest types, and used indicator species analysis to identify species driving the differences between forest types. We then compiled sets of proposed models for predicting plant community structure, and used Akaike's information criterion (AIC_C) to determine the support for each model. Model averaging was used to make multi‐model inferences if no single model was supported. Results Within the lower montane zone, pine–oak forests had greater understorey plant cover, richness and diversity than pure stands of ponderosa pine (Pinus ponderosa P. & C. Lawson var. scopulorum Engelm.). Plant cover was negatively related to time since fire and to ponderosa pine basal area, and was highest on northern slopes and where Gambel oak (Quercus gambelii Nutt.) was present. Species richness was negatively related to time since fire and to ponderosa pine basal area, and was highest on southern slopes and where Gambel oak was present. Annual forb species richness was negatively related to time since fire. Community composition was related to time since fire, pine and oak basal area, and topography. Within subalpine forests, plant cover was negatively related to subalpine fir basal area and amounts of coarse woody debris (CWD), and positively related to Engelmann spruce basal area. Species richness was negatively related to subalpine fir basal area and amounts of CWD, was positively related to Engelmann spruce basal area, and was highest on southern slopes. Community composition was related to spruce, fir and aspen basal areas, amounts of CWD, and topography. Main conclusions In montane forests, low‐intensity surface fire is an important ecological process that maintains understorey communities within the range of natural variability and appears to promote landscape heterogeneity. The presence of Gambel oak was positively associated with high floristic diversity. Therefore management that encourages lightning‐initiated wildfires and Gambel oak production may promote floristic diversity. In subalpine forests, warm southern slopes and areas with low amounts of subalpine fir and CWD were positively associated with high floristic diversity. Therefore the reduction of CWD and forest densities through managed wildfire may promote floristic diversity, although fire use in subalpine forests is inherently more difficult due to intense fire behaviour in dense spruce–fir forests.     

Slow understory redevelopment after clearcutting in high mountain forests

Besides natural tree regeneration itself, the development of the forest understory community is highly indicative of the ecological recovery of forest stands post-harvesting, and therefore of the sustainability of forest management. High mountain forests might show particularly slow recovery of the understory plant community because of harsh environmental conditions. We compared understory community richness and composition among three age classes of forest stands in the subalpine Engelmann Spruce–Subalpine Fir zone in the interior of British Columbia, Canada. Species composition was found to differ significantly between mature stands (>110 years old and never harvested) and both recent clearcuts (5–8 years old) and the oldest clearcuts present in the study area (second growth: 24–28 years old). A non-metric multidimensional scaling (NMDS) ordination revealed no unidirectional return of species composition in harvested stands towards that of mature forest; indeed, plots in recent clearcuts and second growth stands were similar to one another and clearly separated from the mature stands. Indicator Species Analysis revealed that moss species were particularly indicative of mature forest, with four moss species being common in mature stands but absent from both younger stages. Compared to what has been reported for lower elevation coniferous forests, e.g. in the U.S. Pacific Northwest, redevelopment of the understory appears to be slow after harvesting in these high elevation mountain forests. Rotation intervals that consider the natural temporal pattern of species turnover and the occurrence interval of major natural disturbances (here: fire) should provide effective approaches to sustainable forest management of these forests.     

Half a century of succession in a temperate oakwood: from species‐rich community to mesic forest

Aim Lowland woodlands in Europe went through dramatic changes in management in the past century. This article investigates the influence of two key factors, abandonment of coppicing and increased pressure of ungulates, in thermophilous oakwoods. We focused on three interconnected topics: (1) Has the assumed successional trend lead to impoverishment of the vegetation assemblages? (2) Has it resulted in vegetation homogenization? (3) Are the thermophilous oakwoods loosing their original character? Location Czech Republic, Central Europe. Methods The vegetation in 46 semi‐permanent plots was recorded three times: firstly, shortly after the abandonment of coppicing (1953) and then, after four to six decades of secondary succession and strong game impact (1992 and 2006). Overall trends and changes in species spectra were analysed. Results There is a marked successional shift towards species‐poorer communities growing in cooler, moister and nutrient‐richer conditions. The change was significantly different in parts affected and unaffected by high numbers of ungulates yet only for herbs, not the woody species. However, observed change in species composition was not accompanied by significant homogenization process that is the general process reported from elsewhere. A sharp decline in plant species typical for thermophilous woodland communities and in endangered species indicates that the original character of the woodland has been gradually lost. Main conclusions Thermophilous oakwoods have been largely replaced by mesic forests. Lowland oakwoods in continental parts of Europe historically depended on active management, which kept the understorey conditions light and warm. Successional processes in the 20th century caused a critical loss of species diversity at various spatial levels. However, artificially high numbers of ungulates, which otherwise have a negative impact, probably held up succession, so that the changes may still be reversible.