Patterns of functional diversity of two trophic groups after canopy thinning in an abandoned coppice
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Coppice abandonment had negative consequences for the biodiversity of forest vegetation and several groups of invertebrates. Most coppicing restoration studies have focused only on a single trophic level despite the fact that ecosystems are characterized by interactions between trophic levels represented by various groups of organisms. To address the patterns of functional diversity in the perspective of coppicing restoration, we studied the short-term effects of conservation-motivated tree canopy thinning in an abandoned coppice with standards in Central Europe, a region where such attempts have been rare so far. The functional diversity of vascular plants and spiders, chosen as two model trophic groups within the forest ecosystem, was compared between thinned and control forest patches. To characterize functional patterns, we examined several functional traits. These traits were assigned to two contrasting categories: response traits reflecting a change of environment (for both vascular plants and spiders) and effect traits influencing the ecosystem properties (only for vascular plants). Functional diversity was analysed by CCA using two measures: community-weighted means (CWM) and Rao’s quadratic diversity (RaoQ). CCA models revealed that the canopy thinning had a positive effect on the diversity of the response traits of both trophic groups and negatively influenced the diversity of effect traits. In addition, we found distinct seasonal dynamics in functional diversity of the spider communities, which was probably linked to leaf phenology of deciduous trees. We conclude that canopy thinning affected functional diversity across trophic groups during the initial phase of coppicing restoration. With necessary precautions, careful canopy thinning can be effectively applied in the restoration of functional diversity in abandoned coppices.
Keywordscoppice restoration effect traits functional diversity response traits spiders trophic groups vascular plants
Coppicing, an ancient forest management system, was for centuries the prevailing method of forest use in Europe (Buckley and Mills 2015a; Szabó et al. 2015). It can be assumed that a whole range of species and functional groups, as well as types of temperate forest ecosystems, have developed and persisted via traditional management (Kirby and Watkins 2015). Coppice abandonment, a common feature in 20th-century forestry, has led to a decline in the biodiversity of vascular plants and several groups of invertebrates (Beneš et al. 2006; Van Calster et al. 2007; Hédl et al. 2010; Kopecký et al. 2013). Therefore, coppice restoration has been thoroughly debated especially in the United Kingdom (e.g. Buckley 1992; Buckley and Mills 2015b; for critical considerations see Hambler and Speight 1995a,b). Elsewhere, however, the potential consequences of restoring coppicing practices in long-abandoned coppices are largely unknown. Coppicing includes a wide range of tree cutting intensities and in practice can vary from selective cutting to extensive clear-cuts. Careful tree canopy thinning can be a starting point through which the effects of coppicing renewal on biodiversity can be assessed.
The effect of tree canopy thinning on the diversity, composition and functional properties of forest ecosystems has been richly documented (Aubert et al. 2003; Adrian et al. 2009; Verschuyl et al. 2011; Vild et al. 2013). However, the majority of works studying the effects of forest management on biodiversity focused on species diversity (Battles et al. 2001; Dodson et al. 2008; Adrian et al. 2009), while impacts on functional diversity have remained practically unexamined. Understanding functional diversity patterns is important because species diversity is not always the best predictor of ecosystem quality (Tscharntke et al. 2012). Changes in species composition and functional characteristics can have a larger effect on ecosystem functioning than on species richness or diversity (Wilsey and Potvin 2000). Despite the fact that most studies confirm the positive effect of thinning management on understorey species richness, the effect of various intensities of thinning on multiple trophic groups within a given forest ecosystem is still not well understood (Collins et al. 2007; Knapp et al. 2007; Dodson et al. 2008; Atkinson et al. 2015). This lack of knowledge can pose serious limitations because species communities form ecosystems via direct and indirect trophic interactions (Schmitz and Suttle 2001; Ohgushi 2008). Understanding a species’ impact on ecosystem processes (i.e. resistance, resilience and stability) requires a complex approach, hence the inclusion of various trophic groups is of great importance.
Forest canopy thinning is a disturbance process that directly affects many trophic and functional groups by releasing or reducing resources such as light, moisture, living space and nutrients (Kaye and Hart 1998; Gundale et al. 2005). Thus, thinning-induced plant response will influence the diversity, composition and functional properties of other trophic levels, for example of the herbivorous insects and their predators they comprise (Southwood 1977; Price et al. 1980; Pearson and Dyer 2006). Species which will occur in post-logged sites should be filtered from the pool of potential colonist species with an effective competitive ability under specific habitat conditions (Townsend and Hildrew 1994). We assume that species composition will be determined by the physical complexity of the habitat in the sense of the ‘habitat template theory’ (Southwood 1977, 1988; Aarssen and Schamp 2002). Structural and trophic complexity does not increase linearly with succession in temperate regions (Southwood 1977). Vegetation often shows greatest diversity in relatively early stages of secondary succession, before strong competitors start to dominate. Diminishing the overwhelming influence of forest succession by canopy thinning can therefore have a positive effect on the species and functional diversity of certain groups of organisms. We also expect the ‘stress-dominance hypothesis’ will help explain the effects of canopy thinning. According to this hypothesis, the relative importance of environmental filtering increases and competition decreases along the disturbance gradient (Tilman 2004; Coyle et al. 2014). Early stages of succession should be dominated by species with generalist-feeding strategies and broad niches resistant to environmental filtering, as well as pioneer species with the ability to easily disperse and quickly utilize resources but with a weak ability to compete (Pianka 1970; Southwood 1977). In the later phases of ecosystem succession following canopy thinning, the functional properties of a species will mainly reflect competition for resources (Coyle et al. 2014).
The functional traits of species comprising communities are effective measures of various patterns, including properties of successional dynamics. The relative proportions of functional traits can be used to calculate functional diversity (Mason et al. 2005). The concept of functional diversity is a valuable tool for understanding what determines community structure (Gerisch et al. 2012; Kašák et al. 2015). Functional traits can be divided into the two main categories – species effect traits and species response traits (Lavorel and Garnier 2002; Lavorel et al. 2007). Functional effect traits are considered to influence ecosystem properties whereas functional response traits are considered to reflect them (Lavorel and Garnier 2002). For a concrete purpose, the trait groups can be associated with the strategies adopted by individual species to cope with environmental changes such as periodical coppicing. Moreover, the two groups of traits are not completely separable and assignment to one or another trait group is relative, i.e. regarding the context of studied ecosystem. Next to that, traits can be considered according to their effect on ecosystem function, hence traits with strong (assumed effect traits) and weak (assumed response traits) effects (Pakeman 2011). Two main approaches to calculating functional trait diversity exist. The community-weighted mean is the mean trait value weighted by relative species abundance. It gives a dominant trait value in a community. Rao’s quadratic entropy index (RaoQ) quantifies the distribution of trait values among species (Mouchet et al. 2010; Roscher et al. 2012). Communities simultaneously indicating high trait differentiation and high species abundances have high RaoQ values.
The purpose of this study is to evaluate the effect of conservation-motivated tree thinning by comparing functional diversity two different trophic groups: vascular plants and spiders. Our aim is to examine the response patterns and assess appropriateness of thinning intervention for biodiversity conservation. The main hypothesis is that (1) vascular plants and spiders respond similarly to canopy thinning. We assume a pulse release of resources such as light and nitrogen, and increased spatial heterogeneity of these resources because of tree canopy differentiation by selective cutting (see the description of the thinning intervention in Methods). In connection to the resource differentiation, we hypothesize that (2) canopy thinning has a positive effect on functional diversity of response traits and negative influence on functional diversity of effect traits. This prediction reflects the assumption that canopy thinning is associated with traits linked to early successional stages. Corresponding strategies are rapid colonization of suitable patches. Negative influence of canopy thinning corresponds with adaptations to late successional stages with closed canopy. Appropriate traits directly influence environmental conditions. As an additional insight into the short-term functional patterns, we examined spider communities during a vegetation season. We hypothesize that (3) tree leaf shading has a seasonally variable effect on the diversity of functional characteristics of spider communities.
Material and Methods
The study site was Děvín Hill, southern Moravia, Czech Republic (48°52′29.3″ N 16°39′11.9″ E) and has an area of about 380 ha, three-fourth of which are covered with temperate deciduous forest. The site has a temperate climate of subcontinental character with dry periods occurring from late spring to autumn. Average yearly temperatures are 9 to 10°C; precipitation is on average 550 mm·yr−1. Soils here have developed on limestone and loess substrate and are very rich in nutrients; litter decomposes rapidly into a mull-type humus. Our study plots were situated on a slope with a northwesterly aspect and were located 270 to 400 m a.s.l. They are covered with a deciduous forest dominated by broad-leaved lime (Tilia platyphyllos), among other species such as ash (Fraxinus excelsior), three species of maple (Acer plananoides, A. pseudoplanatus, A. campestre), hornbeam (Carpinus betulus), elm (Ulmus glabra) and oak (Quercus petraea). Vegetation is species-rich; about 630 vascular plant species occur on the whole site, and the forest communities can contain up to 70 species per 200 m2 (Nature Conservation Agency of the Czech Republic 2015 and own unpublished results).
The present-day forests on Děvín Hill are among the best examples of abandoned coppices in Central Europe. Most compartments still retain characteristic coppicing features such as coppice stools, some of them apparently centuries old. Historical archival research has demonstrated that nearly the entire forest area of Děvín Hill was managed under a coppicing regime for centuries. The abundance of standard trees largely varied through the site’s history. Currently, mainly oak standards are left in the abandoned coppices. The coppicing management system was exceptionally stable from at least from the fourteenth century until the early twentieth century; for details see Szabó (2010) and Müllerová et al. (2014, 2015). In 1946 the site became a nature reserve with the aim to preserve valuable plant and animal species and communities. Following the then-prevailing conservation paradigm that viewed coppicing as harmful to forest ecosystems, over the past six decades forest management has been based on the principle of least intervention. As a consequence the forests of Děvín have undergone succession from short-rotation coppices less than 40 years old to old-growth forests often over 100 years old (Müllerová et al. 2015). This transformation has resulted in a considerable decline of vegetation biodiversity (Kopecký et al. 2013). The current nature conservation strategy envisages coppicing restoration. The intervention commenced in 2009 with canopy thinning in selected compartments are an attempt to support the vanishing biodiversity.
In all four compartments, the composition of vascular plant and spider communities was sampled using plots and traps, respectively. Sampling followed canopy thinning with a one- to two-year lag and thus reflected the reaction of biotic communities to the relatively immediate effect of thinning. Vascular plants were sampled using circular plots with a radius of 1 m; thus the plot size was 3.14 m2. Five such plots were regularly clumped within squares of 15 by 15 m. Six such clumps and 30 plots were symmetrically placed in the intact forest compartments; six clumps were in thinned compartments comprising another 30 sampling plots. The presence of all vascular plant species in the forest understorey (up to 1.3 m height) was recorded in August 2011. No additional properties of the plant species were recorded during the sampling.
Spiders were sampled using pitfall traps. We used 500 ml plastic cups containing 4 % formaldehyde solution as a fixative medium. A row of three pitfall traps spaced five metres apart was placed in each compartment. The traps were emptied five times during the vegetation season: on 6 June, 2 July, 4 August, 4 September and 2 October 2010. The species and sexes of the collected specimens were determined. The number of individuals per species and sex were recorded. Pitfall traps were used as a standard method to evaluate epigeal spider fauna. Traps were placed on sites during the main vegetation season, between May and October. Using this method we collected nearly the entire spectrum of ground-dwelling spiders, even species that mature during autumn and live during winter time (these usually start to hatch in September). Although only pitfall traps aimed at epigeal species were used, tree species were also caught. In each compartment the same trap spacing and timing was used. Pitfall traps are able to produce a representative spider sample, also with the most ecologically important species (e.g. Niedobová and Fric 2014).
Functional traits used in the analysis. Sources of trait values for vascular plants are Ellenberg indicator values (EIV), BiolFlor (BF) and LEDA (L). Sources of trait values for spiders are Buchar and Ruzicka (2002), and Kasal and Kaláb (2015). Expected change in the functional traits values following canopy thinning is indicated.
Functional trait type
Light affinity (EIV)
Moisture affinity (EIV)
Soil seed bank (L)
Increase (more persistent)
Specific leaf area (L)
Plant height (L)
Leaf phenophase (BF)
Vascular plant traits were obtained from three resources: Ellenberg indicator values (Ellenberg et al. 1992), LEDA (Kleyer et al. 2008) and BiolFlor (Klotz et al. 2002). Plant traits classified as effect traits were specific leaf area, plant height and leaf phenophase, while traits considered as response traits were light affinity, moisture affinity and persistent soil seed bank. For an explanation and quantification of the particular traits, see the cited database resources. The selected effect traits reflect plant strategies to cope with increasing competition during the forest succession, while constituting the properties of environment at the same time. Taller plant species will tend to dominate in succession, and species with greater leaf area would become more abundant in increasingly shadier conditions. Both traits will have an effect on environmental properties experienced by organisms, including vascular plants and spiders. The latter group would respond to variation in leaf phenophase because foliage development causes changes in shading during the vegetation season. The selected response traits help to quantify the reaction of plant species on changing amount and heterogeneity of important resources, which is obvious for light and moisture affinity. Soil seed bank in periodically thinned forests would change to more persistent because shifting environment would require persistent seeds to survive of unfavourable conditions.
For spiders, only response traits were considered plausible. These traits included light affinity, humidity affinity and stratum affinity. They are based on species environmental requirements described by Buchar and Ruzicka (2002), who established them using catalogue data from the Czech Republic (more than 150,000 records). Similar categories were also established by Kasal and Kaláb (2015). We used species affinity to humidity (very dry, dry, semi-humid, humid and very humid), light (open, semi-open, partly shaded, shaded and dark habitats) and forest vertical stratum (underground, ground layer, vertical surfaces, herb layer, shrub layer, tree trunks and canopies). The affinities of species to trait values, ranging from 1 to 5, were determined subjectively. For the expected changes in individual plant and spider traits following the canopy thinning, see Table 1.
A total of 36 species of spiders (302 specimens) and 86 species of vascular plants were analysed statistically. We used canonical correspondence analysis (CCA) in the statistical software Canoco 5 (ter Braak and Šmilauer 2012) to test the effect of canopy thinning on species composition and abundances. Vascular plant species data were obtained by determining the presence of individual species in plots; spider species data were acquired from pooled abundances across sexes. We chose presence or absence of canopy thinning as explanatory variable. Species abundance of spiders was transformed by decimal logarithm. Calculated P-values were adjusted for multiple testing using Bonferroni correction. The significance of the canonical axis was tested with the Monte-Carlo permutation test (5,000 permutations). To achieve correct Type I error estimates, the permutation test was restricted by using a hierarchical design. The effect of canopy thinning on vascular plants was analysed with a permutation test defined by 15 split-plots in each of the 4 whole plots, e.g. forest stands with thinning treatment. Statistical testing was by permutation of plots independently across the whole plots while split-plots were held together without permutation. For the analysis of spiders, we had to arrange a different statistical design following the sampling design. The permutation test used 10 split-plots (using data from two sampling years) within each of 4 whole plots – forest stands with thinning treatment. In addition, for analysing spiders we restricted the permutation to a time series because of autocorrelation between individual observations. The cases were arranged along a time scale, and therefore during permutations we had to respect the autocorrelation structure that bands the explanatory and response data. Therefore whole plots were freely exchangeable and split-plots were restricted for time series.
Functional diversity was calculated as RaoQ separately for spider and for herbaceous functional and effect traits. RaoQ is defined as the sum of pairwise distances between species divided by relative abundance and is largest when species with large trait differences reach similar high abundances (Mouchet et al. 2010). In addition, we calculated community-weighted means (CWM) of trait values to show how the particular species traits are related to canopy thinning. CWMs were than passively projected as functional traits in the ordination diagrams. The CWM is calculated by dividing mean trait values by species relative abundance indicating the dominant trait value in the community (Roscher et al. 2012). We used CWMs of functional traits as explanatory variables to find out which of them explain significant variability in species data. A CCA model with the same restricted-permutation design was used to analyse the CWMs of functional traits. Forward selection was then applied to select significant CWM traits. These traits and the functional diversity were then passively projected to the ordination space.
The relationship between the functional diversity of spiders and seasonality was fitted by generalized additive model (GAM) with Gaussian error distribution. We used an F-test to determine the significance of the relationship. To reveal if the species composition of the five subsequent samplings really differed we used analysis of similarities (ANOSIM, part of the ‘vegan’ package in R statistical software). ANOSIM is based on computing the rank order of dissimilarity values within and between groups. A dissimilarity matrix was calculated from the species matrix using Bray-Curtis distance. To assess the significance of the ANOSIM statistic, a permutation test was used (1,000 permutations). If two groups of samplings really differ in their species composition, the ANOSIM statistic R will be greater than zero. Thus, compositional dissimilarities are always greater between groups than within groups.
Overall Effects of Canopy Thinning
Response of Vascular Plant Functional Diversity
Significance of spiders and herbaceous plants functional traits used as explanatory variables in the forward selection for the CCA models.
Functional trait type
Model variability explained [%]
Functional response traits
Soil seed bank
Functional effect traits
Specific leaf area
Functional response traits
Response of Spider Functional Diversity
Seasonal Variability of Sider Functional Diversity
Reactions of Trophic Groups to Canopy Thinning
Our results indicate that forest canopy thinning influenced compositional patterns and had positive effects on functional diversity of the two simultaneously trophic groups under study – vascular plants and spiders. The latter aspect is relatively novel, and considering the coppicing restoration, our study appears to be unique. The findings are generally consistent with those of the meta-analysis by Verschuyl et al. (2011) showing that forest thinning has a mostly positive effect, or at most a neutral impact, on the biodiversity of various animal groups. Coppicing as a system using frequent canopy thinning is sometimes sharply criticized as being harmful to biodiversity and functioning of forest ecosystems (e.g. Hambler and Speight 1995b). Our results led us to a different conclusion, but it should be considered that it is related only to an early development immediately following coppicing of modest intensity. The positive effect of canopy thinning on biodiversity in the initial years of succession has been however documented by several studies (e.g. Collins and Pickett 1988; Wayman and North 2007; Verschuyl et al. 2011). Subsequent development can vary greatly, but in general biodiversity likely declines as the canopy closes. In order to make conclusions on the development of restored coppices that are valid for periods longer than a few years after tree cutting, greater observation periods are required (e.g. Ash and Barkham 1976; Mason and MacDonald 2002). Contrasting patterns when distinguished for response and effect traits in vascular plants may be explained by differing strategies adopted by species in relation to early versus late succession stages. The positive effects the spatial heterogeneity created by canopy thinning had on spider functional diversity was further supplemented by the similar effects of temporal heterogeneity due to tree leaf development throughout the growing season. However, vascular plants and spiders are reasonably distinct trophic groups, so their reactions to canopy thinning are further discussed separately.
Response of Vascular Plants
Understorey vegetation shows a physiologically-established response to succession (Bazzaz 1979). Our results suggest that increased light was the critical resource influencing species composition. This factor is important for the overall performance of forest herb plants (Whigham 2004). Many studies have shown that canopy openness, regardless if it is the result of natural processes or intentional thinning like coppicing, has a significant, complex effect on understorey herb species performance (e.g. Goldblum 1997; Valverde and Silvertown 1998; Mason and MacDonald 2002; Lindh 2008; Van Calster et al. 2008; Scanga 2014), demonstrating the positive effect of the increased light availability on the diversity of forest understorey vegetation (e.g. Ares et al. 2009; Bailey et al. 1998; Whigham 2004). Other studies, however, determined immediate negative effects of canopy thinning on understorey vegetation (e.g. Adrian et al. 2009). It was postulated that disturbance can initially homogenize vegetation structure and increase light uniformity (Beggs 2005; Bartemucci et al. 2006). We argue that the intensity and extent of canopy thinning plays a critical role. Intensive disturbance could affect the diversity at the species and functional levels by reducing the number of disturbance-sensitive taxa (e.g. Griffis et al. 2001). On the other hand, thinning not overly destructive to the physical environment can support immigration and the establishment of populations of species benefiting from the release of light and nutrient resources.
When partitioning the functional diversity for response and effect traits, we identified two complementary stories. Not surprisingly, tree thinning was positively associated with the diversity of plant adaptations responding to environmental changes, and negatively with adaptations constituting stable environment (McLachlan and Bazely 2001; Wellnitz and Poff 2001; Decocq et al. 2004). Coppicing restoration positively influenced occurrence of species with strategies suitable to early successional phases, which were mostly associated with response traits. Persistent soil seed bank or high light affinity is shared by species preferring frequently disturbed habitats. The occurrence of a number of typical open-habitat species supports this interpretation. They are largely pioneer species that rapidly colonize the newly open patches following canopy removal (Ash and Barkham 1976; Ohtsuka et al. 1993). In the presented study, this group includes Moehringia trinervia, Taraxacum sect. Ruderalia, Trifolium repens, Ajuga reptans, Urtica dioica, Scrophularia nodosa and Sambucus nigra; for an additional list of species emerging during the coppicing restoration in a secondary forest, see Hédl et al. (this issue). On the other hand, undisturbed forest sustains the diversity of species sensitive to disturbances, mainly shade-tolerant competitors (Southwood 1977; Bazzaz 1979). They are characterized with adaptations as tall growth and high moisture affinity, which is associated with high specific leaf area. Several shade-tolerant species were associated with our control plots, e.g. Aconitum lycoctonum, Asarum europaeum, Chaerophyllum temulum or Viola mirabilis. Some of them can significantly influence the species composition of the understorey communities (cf. effect traits).
Response of Spiders
We expected that shifts in particular functional traits of vascular plants (namely specific leaf area, plant height, and leaf phenophase) would be reflected in changes in spider species composition and the diversity of spider response traits. Indeed, the CCA model revealed that spider functional diversity showed the same trend as understorey vegetation. In addition, we assumed that the greater variability of functional groups of spiders on the thinned sites could be caused by the increased spatial and temporal heterogeneity of physical conditions, including the vegetation structure (Southwood 1977, 1988; Schuldt et al. 2012; Mori et al. 2015). The hypothesis predicting higher spatial heterogeneity after canopy thinning can be partially supported by our results. A positive effect of thinning on vertical stratification of spider habitats was revealed. We therefore assume that the thinned patches supported spiders favouring the tree canopy (Anyphaena accentuata, Tegenaria ferruginea, Segestria senoculata) because these spiders moved from tree canopies to shrub canopies.
Moreover, larger plant size was also positively correlated with canopy thinning. This relationship implies that after canopy thinning high light availability favoured larger-sized plant species. We can conclude that vegetation structure complexity is also an important factor positively influencing species richness (Schuldt et al. 2012; McDonald 2015). It was demonstrated that species number may be related to foliage height diversity and cover (Roth 1976). The non-linear trend between succession and spatial complexity can thus lead to a great diversity of spiders and plants after canopy thinning (Southwood 1977).
The random colonization of species with unique combinations of functional traits is a typical phenomenon for early succession stages supporting functional trait diversity (e.g. Kašák et al. 2013; Hodeček et al. 2015). The lower functional diversity of spiders in control patches can be explained by the strong shading impact of trees and therefore the relative environmental uniformity of these patches (Beggs 2005; Bartemucci et al. 2006). Our results support this prediction because none of the functional traits for spiders show a positive correlation with control patches. Similar results are also known for other groups of epigeal invertebrates as well as spiders (Spitzer et al. 2008; Purchart et al. 2013).
The development of spider functional diversity during the growing season showed a significant unimodal pattern. We explain this observation by tree leaf phenology, which influences the temperature and light heterogeneity of the forest throughout the year (Anderson 1964; Hutchison and Matt 1977). The high heterogeneity of the physical factors at the beginning and end of the growing season leads to a high level of functional diversity. Interestingly, the increases in functional diversity were not caused by species migration within the habitat, but by the colonization of the habitat by new species from the surrounding areas.
Our study shows that careful canopy thinning in an abandoned coppice can have an immediate effect on the functional diversity of species belonging to different trophic groups. Whether functional diversity responds positively or negatively to canopy thinning depends on the functional trait type (i.e. response and effect traits). The functional vascular plant diversity was largely associated with adaptations to light and moisture availability, while spider diversity was positively linked to the light and vegetation structure, and changed during the vegetation season with the development of tree leaves. Canopy thinning can be generally regarded as a way towards ecosystem-friendly coppicing restoration. Precautions need to be taken for various trophic groups, trophic conditions and thinning intensity. We therefore suggest that future research determine the habitat-differentiated intensity of canopy thinning in order to ensure optimal ecosystem stability and functioning.
The findings published in this paper were obtained though financial support from the Grant Agency of the Academy of Sciences of the Czech Republic, grant AV0 IAA600050812 ‘Lowland woodland in the perspective of historical development’, and from the Ministry of Education, Youth and Sports of the Czech Republic, grant CZ.1.07/2.3.00/20.0267 ‘Coppice forests as the production and biological alternative for the future’. Additional support came from the European Research Council, (FP7/2007-2013) grant 278065 ‘Long-term woodland dynamics in Central Europe: from estimations to a realistic model’, from the project MUNI/A/1048/2015, and from the Czech Academy of Sciences, long-term research development project no. RVO 67985939.
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