Changes of Forest Abiotic Environment in the Western Carpathians Assessed Using Phytoindication

JOURNAL OF FOREST SCIENCE, 60, 2014 (4): 133–142

Changes of forest abiotic environment in the Western Carpathians assessed using phytoindication

W. Żelazny

Department of Forest Protection and Wildlife Management, Faculty of Forestry and Wood Technology, Mendel University in Brno, Brno, Czech Republic

ABSTRACT: To avoid ecosystem degradation, forestry planning needs to be based on current information about the state of forest environment. Phytoindication is an inexpensive tool that allows tracking the environmental change at fine spatial scales. The present study uses this approach to assess changes within abiotic conditions of forests in the area of the Moravian-Silesian Beskids Mts. (Czech Republic). Phytosociological relevés collected in 2013 at 118 permanent plots were compared with records from the 1960s and 1970s. The changes were expressed using average Ellenberg’s Indicator Values and units of the Czech Forest Typological System. Persisting soil acidification was detected, and linked to industrial deposition and planting of Norway spruce beyond its natural range. Conversion towards a higher share of broadleaved species was suggested to support soil recovery.

Keywords: Beskids Mts.; Ellenberg’s indicator values; forest typology; Norway spruce; soil acidification

The Carpathian Mountains belong to the most nent environmental conditions” (Viewegh et al. important European biocentres, being a provider 2003). It can be argued, however, that these charac- of ecosystem services on which many human com- teristics are not only variable but given the current munities depend (Gurung et al. 2009). Among scale of anthropogenic disturbance (Steffen et al. the services delivered by mountain forests, there is 2011) while rate of their change has dramatically the supply of wood and other forest products. By accelerated. The increasing speed of environmental careful planning, modern forestry strives to opti- change calls for an introduction of adaptive man- mize these outputs, at the same time avoiding the agement strategies into forestry practice. In order ecosystem degradation. Management plans have to to be successful, this approach needs to be based consider the parameters of the abiotic environment on current data (Lindner et al. 2010). of each forest stand (Vacek, Balcar 2004). The goal of the present study is to analyze recent In the forestry practice of the Czech Republic, changes within the forest abiotic environment of a these characteristics are expressed using units of Western Carpathian mountain range using phyto- the Czech Forest Typological System. Among the indication. The trends of change are described by units there are forest vegetation zones which take referring to average Ellenberg’s Indicator Values their names from edificator species and describe (AIVs) and the units of the Czech Forest Typologi- the variability of local climate with changing alti- cal System. Recommendations for the local forest tude and exposition, and edaphic categories which management are given. describe the soil trophic and hydric regime and are According to Hédl (2004), three major trends denoted using single-letter mnemonics (Viewegh of anthropogenic change may currently be tak- et al. 2003). ing place within the abiotic environment of forest The parameters of the forest abiotic environment ecosystems of Central Europe: (1) soil acidification covered by these units are referred to as “perma- caused by the deposition of air pollutants; (2) eu-

Supported by the Mendel University in Brno, IGA LDF Grant No. 23/2013.

J. FOR. SCI., 60, 2014 (4): 133–142 133 trophication due to the pollution and the overpop- tained values did not suggest high acidification lev- ulation of herbivores; (3) shifts related to the global els (Staszewski et al. 2012). climate change. The data on climatic changes are available in the With the exception of minor fragments, the Car- form of meteorological reports and the outputs of pathian Mts. lie within the boundaries of the former climatological models (e.g. Nogués-Bravo et al. Communist Bloc, where the economy was based 2007; Hlásny et al. 2011). According to the pre- on heavy industry, and environmental regulations dictions, by the end of the 21st century, lower Car- were sloppy. For decades, the ecosystems were ex- pathian ranges are going to be subjected to drought posed to high air pollution levels, resulting in the events, whereas at higher elevations temperature deposition of sulphur and nitrogen compounds, increases are likely to occur ‒ potentially boosting which have been contributing to the decline of forest productivity (Lindner et al. 2010; Hlásny forest health. The affected stands include Norway et al. 2011). Unfortunately, due to the coarse spatial spruce (Picea abies [L.] Karsten) monocultures, scales involved in such assessments (Nogués-Bra- whose widespread presence has been the legacy of vo et al. 2007), the possibilities of their application the 19th century forestry policy (Main-Knorn et in adaptive forest management are limited. This is al. 2009). especially true of mountain ranges, which are char- After the Fall of Communism, the industrial emis- acterized by rich relief, resulting in a high variation sions have been reduced and many forest stands of habitat types (Nepal, Chipeniuk 2005). have been undergoing conversion toward a higher A particular approach to environmental monitor- share of broadleaved species (Main-Knorn et al. ing that does not rely on specialist equipment and 2009). While in theory both trends create an op- expensive analyses (Diekmann 2003) and which portunity for the edaphic environment to recover, makes it possible to obtain data with high spatial evidence of such a recovery has been scarce so far: resolution (Balkovič et al. 2010) is to use phyto- recent measurements of the forest environment indication. Under this approach repeated sampling quality in the Czech (Novotný et al. 2008), Slovak of phytosociological permanent plots is performed, (Sitková et al. 2010) and Romanian (Bytnero- and observed shifts in the vegetation composition wicz et al. 2005; Badea et al. 2012) parts of the are interpreted in terms of environmental change Carpathian Mountains give a picture of high soil by referring to the changes of average Ellenberg’s acidification levels that have been persisting despite Indicator Values (AIVs). a substantial reduction of pollutant concentrations The reports from phytosociological studies car- in the atmosphere. The affected ecosystems include ried out in the Czech (Šamonil, Vrška 2007) and not only those located in the proximity of pollution Polish (Durak 2010, 2011) Carpathians mention sources: Šebesta et al. (2011) studied changes that a decrease of AIV R, indicating a trend towards had occurred since the 1930s in the soil environ- the lower soil pH or calcium concentration (Diek- ment of a primeval forest ecosystem in the Ukrai- mann 2003). An opposite direction of change was nian Carpathians, close to the Romanian border. detected at Norway spruce-dominated stands in Despite the large distance dividing their study area the Ukrainian part of the mountains; interestingly, from industrial centres, the evidence of acidifica- the result was not consistent with the results of tion was present in the samples. The results agree soil analyses (Šebesta et al. 2011). According to with the findings of Oulehle et al. (2010) per- the authors, this discrepancy could be explained taining to the same mountain range. Interestingly, by high resistance of the phytocoenoses to habitat the latter authors did not observe substantial soil disturbances or by colonization of low tree layers acidification in another part of the Ukrainian Car- by broadleaved woody species. The same study re- pathians, next to the Slovak border. This discrep- ported increasing AIV N, what can be interpreted ancy was explained as the result of differing par- as soil enrichment in nitrogen, or more intensive ent bedrocks at both localities. On the other hand, biomass production (Diekmann 2003). The au- the analysis of the soil drainage water collected in thors linked this observation to atmospheric nitro- the area less affected by acidification revealed in- gen deposition and shifting forest developmental tensive leaching of nitrogen, suggesting a high level phases (Šebesta et al. 2011). A different result was of eutrophication (Oulehle et al. 2010). Another obtained by Durak (2011) in the Polish Carpath- notable exception is the Polish study on the forest ians. The AIV N decrease was interpreted as the health status in national parks. On several perma- consequence of silvicultural operations. Some re- nent plots, located mostly in the Carpathians, the searchers (Šamonil, Vrška 2007; Durak 2010; authors surveyed Ca to Al ratios of soil. The ob- Šebesta et al. 2011) reported changes of AIV L,

134 J. FOR. SCI., 60, 2014 (4): 133–142 but as pointed out by Diekmann (2003), the reli- whereas the remaining fraction showed the oppo- ability of this light-related parameter is low, as its site trend of migration. values do not correlate well with measured light intensity over short gradients. Owing to the fact that the definition of the units of MATERIAL AND METHODS the Czech Forest Typological System is closely related to the distribution of potential vegetation (Viewegh The research was conducted at the top elevations of et al. 2003), they can be used in a similar way like the Moravian-Silesian Beskids Mts., which belong to AIVs to monitor changes within the forest abiotic the Western Carpathians and are located in the east- environment. Viewegh (1999) used this approach to ern part of the Czech Republic. This mountain range interpret vegetation shifts in the Moravian-Silesian is characterized by a high share of forest cover, reach- Beskids Mts. (Western Carpathians). Between 1984 ing 75% of the total land area, and plant species rich- and 1986, the author resampled 390 permanent plots ness exceeding one thousand taxa. Notable are high and compared the relevés with earlier records, proba- precipitation sums, which amount to 900–1,377 mm bly collected in the 1930s, using an unconstrained or- per annum. Annual mean temperatures are in the dination. By referring to the typological classification 2.3–7.8°C range. The dominating soil type is Cam- of the plots, he concluded that the abiotic character- bisol, which has been formed on the flysch bedrock. istics of the ecosystems had been changing towards The potential vegetation of the area consists mainly of conditions typical of sparse high-mountain stands. European beech (Fagus sylvatica L.) forests, but the The phenomena that are related to the global species has been largely replaced by Norway spruce, climate change and which can be monitored us- planted beyond its natural range and currently pre- ing phytoindication methods include the shifts vailing in the tree composition (Vacek 2003). of mountain vegetation zones towards peak el- Between July and September 2013, 198 phytoso- evations. Under the assumption that the ranges of ciological relevés (hereafter, 2013 relevés) were col- many plant species are constrained primarily by cli- lected at permanent plots maintained by the For- matic factors, with the climate change progression est Management Institute in Brandýs nad Labem, it is reasonable to expect widespread migrations of Czech Republic. The plots selected for the study populations following their shifting niches (Bertin had 400–500 m² surface area and were located in 2008). To my knowledge, no such response of phy- the fir-beech (500–1,000 m a.s.l. altitude range), tocoenoses to the changing environment has been spruce-beech (555–1,120 m a.s.l.) and beech- observed in the Carpathians so far; there are, how- spruce (830–1,220 m a.s.l.) forest vegetation zones. ever, research reports suggesting migrations of this The plots were relocated using a GPS device. At kind have been taking place in some other moun- each locality, the abundance-dominance of vascular tain ranges (e.g. Kelly, Goulden 2008; Bai et al. plant species and the Sphagnum mosses were regis- 2011; Feeley et al. 2011). In the European context, tered on the 11-point Zlatník scale. To reduce the notable is the study by Lenoir et al. (2008), who influence of relocation errors, the relevés collected compared vegetation records that had been collect- in heterogeneous stands were excluded from fur- ed over two time periods in six mountain ranges ther analysis. Due to their insufficient amount, also of Western Europe. The authors concentrated on the data from nature reserves and other small-scale changes within the species altitudinal optima and specially protected areas were discarded, resulting reported that two-thirds of the taxa had changed in the final number of 118 relevés (one relevé per their distribution ranges toward higher elevations, one permanent plot).

Table 1. Typological classification of permanent plots, absolute frequencies given (n = 118)

Edaphic category Forest vegetation zone J D U B A F H S K N Z Y V G O T Beech-spruce 0 0 0 0 0 0 0 1 0 0 2 0 0 0 0 1 Spruce-beech 0 0 0 0 0 6 0 3 0 2 0 0 1 2 4 0 Fir-beech 4 3 2 12 10 24 4 21 9 2 0 1 2 0 2 0

J – eutrophic talus, D – eutrophic colluvial, U – ravines and gulleys, B – typical mesotrophic, A – eutrophic stony-colluvial, F – mesotrophic stony, H – mesotrophic loamy, S – transitional meso- to oligotrophic, K – typical oligotrophic, N – oligotrophic stony, Z – scrub, Y – extreme skeletal, V – eutrophic moist to wet, G – mesotrophic wet, O – mesotrophic gleyed, T – oligotrophic gleyed

J. FOR. SCI., 60, 2014 (4): 133–142 135 The plots show an unbalanced representation vegetated with trees during their resampling at a of typological units (Table 1), reflecting the vari- later time. able frequency of forest site type occurrence in the Considering the limited bioindicative value of mountain range. Among the edaphic categories, tree species found in production forests, the subse- the most common are: the category F, indicating quent parts of the study concentrated on the moss, mesotrophic forests growing on stony slopes, and herb and low shrub layers. The Zlatník abundance- the transitional meso- to oligotrophic category S dominance values were converted into cover per- (Viewegh et al. 2003). Lower is the representation centages and the species that had posed identifi- of typical meso- and oligotrophic forests classified cation problems were amalgamated. Because the into the categories B and K, respectively, as well as relevés had been collected by various persons, and eutrophic stands bound to areas of colluvial nutri- because the taxa in the dataset occurred with vari- ent accumulation (category A). Scarce are the data able frequency, a Wisconsin double standardiza- from forests growing on wet, gleyed soils (catego- tion was applied to the values. ries V, G, O, T) or in extreme environmental condi- The dimensionality of the dataset was then re- tions (categories Z and Y). For a more comprehen- duced to four dimensions using the Nonmetric sive description of edaphic categories the reader Multidimensional Scaling (NMDS) method. The is referred to Viewegh et al. (2003). As far as the ordination was based on the Bray-Curtis distance forest vegetation zones are concerned, the relevés matrix and involved 100 random starting configu- from the fir-beech zone form the bulk of the data- rations. A two-dimensional ordination was also set, whereas the records from higher elevations are tried, but it had to be discarded due to high stress limited in number. value. The ordination plots obtained from the more The data collected at the plots were subjected complex model were partitioned into sub-plots to numerical analysis using the R environment representing units of the Czech Forest Typological (R Foundation for Statistical Computing, Vienna, System. The relevés were assigned to the ordination Austria). In order to describe the changes that had sub-plots according to the typological classification taken place in the vegetation patterns during the of permanent plots provided by the Forest Manage- previous decades, the 2013 dataset was merged ment Institute. with 78 records from the 1960s and 78 records from In the next step, six unweighted average Ellen- the 1970s (hereafter, historical relevés; Fig. 1) col- berg’s Indicator Values (AIV) were calculated for lected at the same permanent plots. The 1960s can each relevé and used as input data for general addi- be characterized as the decade of a sharp increase tive models (GAM). The values fitted by the models of industrial emissions (Oulehle et al. 2010), but were represented graphically in the NMDS ordina- at that time the pollution impact on forest health in tion space using isolines. Additionally, a series of the Moravian-Silesian Beskids area was just begin- mixed-effect regression models, with each AIV as ning to show. The damage culminated at the end of the dependent variable, time as the fixed indepen- the 1970s (Vacek 2003). dent variable and permanent plot as the random The majority of the relevés in the merged dataset intercept variable, was fitted. As some relevés were represent stands where Norway spruce was either missing one or more AIV values, this calculation the dominant or the only tree species at the time of was based on a reduced dataset representing 101 the sampling (Table 2). They are followed by forests plots. The model performance was diagnosed by characterized by a high share of European beech. analysing the distribution of residuals, and its out- Other recorded tree species include: sycamore ma- put was used to estimate 95% confidence intervals ple (Acer pseudoplatanus L.), silver fir Abies( alba of AIV changes over the period between the date of Mill.), Scots pine (Pinus sylvestris L.), and Euro- the oldest relevé (i.e. the year 1961) and the sum- pean larch (Larix decidua Mill.). Some of the per- mer 2013. The linear modelling was repeated using manent plots with high canopy closure at the time weighted instead of unweighted AIVs as dependent of their establishment occurred to be only sparsely variables.

196 1111444444444444444444444444444444444444444444444444444444444444 196 88888888888888 197 222222222222233333333333333333333333333333333333334444444444444444 197 888888888888

Fig. 1. Stem and leaf plot of the years when the historical relevés were collected

136 J. FOR. SCI., 60, 2014 (4): 133–142 Table 2. Tree compositions of permanent plots in different periods of vegetation sampling, absolute frequencies given

Sampling period

Tree composition 1961–1968 1972–1978 2013 (n = 78) (n = 78) (n = 118)

Norway spruce 29 27 34 Single-species, consisting of European beech 4 6 6 Sycamore maple 2 2 0 Norway spruce 25 22 45 European beech 8 13 21 Mixed, dominated by Sycamore maple 4 3 2 Silver fir 5 2 0 Scots pine 1 0 1 With canopy closure not exceeding 10% 0 3 9

The directions of change within the vegetation were translated into changes of the forest abiotic composition of the permanent plots were then as- environment using the results of AIV modelling sessed by comparing the locations of the 2013 rele- and the information about the typological classi- vés in the ordination space in respect to the loca- fication of the plots. As interpretable patterns of tions of the historical relevés. The observed shifts change were visible only in the first two dimensions

(a) J (b) D (c) U (d) B 1.5 1.5 1.5 1.5

1 1 1 1

0.5 0.5 0.5 0.5 2

NM D –1 .5 –1 – 0.5 0 0.511.5 –1 .5 –1 – 0.5 0 0.511.5 –1 .5 –1 – 0.5 0 0.511.5 –1 .5 –1 –0 .5 0 0.511.5 – 0.5 –0 .5 –0 .5 – 0.5

–1 –1 –1 –1

–1 .5 –1 .5 –1 .5 –1 .5

(e) A (f) F (g) H (h) S 1.5 1.5 1.5 1.5

1 1 1 1

0.5 0.5 0.5 0.5 2 S

0 0

NM D –1 .5 –1 – 0.5 0 0.511.5 –1 .5 –1 – 0.5 0 0.511.5 –1 .5 –1 –0 .5 0 0.511.5 –1 .5 –1 –0 .5 0 0.511.5 – 0.5 – 0.5 –0 .5 –0 .5

–1 –1 –1 –1

–1 .5 –1 .5 –1 .5 –1 .5

(i) K (j) N (k) Z (l) Y 1.5 1.5 1.5 1.5

1 1 1 1

0.5 0.5 0.5 0.5 2

NM D –1 .5 –1 – 0.5 0 0.511.5 –1 .5 –1 – 0.5 0 0.511.5 –1 .5 –1 – 0.5 0 0.511.5 –1 .5 –1 – 0.5 0 0.511.5 – 0.5 – 0.5 – 0.5 –0 .5

–1 –1 –1 –1

–1 .5 –1 .5 –1 .5 –1 .5

(m) V (n) G (o) O (p) T 1.5 1.5 1.5 1.5

1 1 1 1

0.5 0.5 0.5 0.5 2

NM D –1 .5 –1 – 0.5 0 0.511.5 –1 .5 –1 – 0.5 0 0.511.5 –1 .5 –1 – 0.5 0 0.511.5 –1 .5 –1 – 0 . 5 0 0.511.5 –0 .5 –0 .5 –0 .5 –0 .5

–1 –1 –1 –1

–1 .5 –1 .5 –1 .5 –1 .5

NMDS1 NMDS1 NMDS1 NMDS1 1960s 1970s 2013

Fig. 2. The first two-dimensions of the NMDS ordination space with observations partitioned into edaphic categories and differentiated according to the sampling period; large symbols represent weight centres of each group

J. FOR. SCI., 60, 2014 (4): 133–142 137

Page 1 J D U B 1.5 1.5 1.5 1.5

1 1 1 1

0.5 0.5 0.5 0.5 2

NM D –1 .5 –1 – 0.5 0 0.511.5 –1 .5 –1 – 0.5 0 0.511.5 –1 .5 –1 – 0.5 0 0.511.5 –1 .5 –1 –0 .5 0 0.511.5 – 0.5 –0 .5 –0 .5 – 0.5

–1 –1 –1 –1

–1 .5 –1 .5 –1 .5 –1 .5

A F H S 1.5 1.5 1.5 1.5

1 1 1 1

0.5 0.5 0.5 0.5 2 S

0 0

NM D –1 .5 –1 – 0.5 0 0.511.5 –1 .5 –1 – 0.5 0 0.511.5 –1 .5 –1 –0 .5 0 0.511.5 –1 .5 –1 –0 .5 0 0.511.5 – 0.5 – 0.5 –0 .5 –0 .5

–1 –1 –1 –1

–1 .5 –1 .5 –1 .5 –1 .5

K N Z Y 1.5 1.5 1.5 1.5

1 1 1 1

0.5 0.5 0.5 0.5 2

NM D –1 .5 –1 – 0.5 0 0.511.5 –1 .5 –1 – 0.5 0 0.511.5 –1 .5 –1 – 0.5 0 0.511.5 –1 .5 –1 – 0.5 0 0.511.5 – 0.5 – 0.5 – 0.5 –0 .5

–1 –1 –1 –1

–1 .5 –1 .5 –1 .5 –1 .5 Fig. 3

D V G O T U 1.5 1.5 1.5 1.5 S B (a) Fi 1r-beech zone (b)1 Spruce -beech zone 1 (c) Beech-spruce zone1 O 0.5 0.5 0.5 0.5 2

S S 1.5 1.5 1.5

F NM D –1 .5 –1 – 0.5 0 0.511.5 –1 .5 –1 – 0.5 0 0.511.5 –1 .5 –1 – 0.5 0 0.511.5 –1 .5 –1 – 0 . 5 0 0.511.5 B –0 .5 1 –0 .5 1 –0 .5 1 –0 .5 S – – – –1 2 1 0.5 1 0.5 1 0.5 H S

–1 .5 –1 .5 –1 .5 –1 .5 A A NMDS1 0 NMDS1 NMDS1 NMDS1 NM D –1.5 –1 –0.5 0 0.5 1 1.5 –1.5 –1 –0.5 0 0.5 1 1.5 –1.5 –1 –0.5 0 0.5 1 1.5 S – – – A 0.5 0.5 0.5 F – – – S 1 1 1 S –1.5 –1.5 –1.5 S S NMDS1 NMDS1 NMDS1 S S 1960s 1970s 2013 F S Fig. 3. The first two dimensions of the NMDS ordination space with observations partitioned forest vegetation zones and differentiated according to the sampling period (a–c); large symbols represent weight centres of each group

of the ordination, the remaining two are not pre- ordination space is occupied by observations origi- sented in this paper. nating from eutrophic sites (category J) and the sites belonging to the mesotrophic category B. In the central part, relevés collected in meso- to oligotrophic (S) RESULTS habitats predominate. These two clusters share the space with the categories A and F, which are assigned The obtained NMDS ordination is characterized to forests growing on slopes. The right-hand side by the stress value close to 0.16, indicating a good fit groups relevés from oligotrophic sites (category K) and an improvement over the two-dimensional ordi- and mesotrophic sites with gleyed soils (category O). nation (stress value of 0.26). After partitioning into The remaining edaphic categories covered by the edaphic categories (Fig. 2), several clusters of histori- study are poorly represented. cal relevés (marked with circles and triangles on the Partitioning according to the forest vegetation

plots) can be distinguished: the left-hand side of the Pagezones 1 (Fig. 3) reveals no apparent altitudinal gradi-

(a) L (b) R (c) N 0.848 0.848 0.848 0.68 0.68 0.68 0.512 0.512 0.512 0.344 0.344 0.344 2

S 0.176 0.176 0.176 D 0.008 0.008Page 1 0.008 M – 0.16 – 0.16 – 0.16 N – 0.328 – 0.328 – 0.328 – – –

0.496 0.496 0.496 – 0.664 – 0.664 – 0.664 – 0.832 – 0.832 – 0.832 –1 – 1 –1 1 1 8 8 8 1 – 08 76 44 12 48 08 76 44 12 48 08 76 44 12 48 – – 0.16 0. 6 0.16 0. 6 0.16 0. 6 0.832 0.664 0.496 0.328 0.832 0.664 0.496 0.328 0.832 0.664 0.496 0.328 0. 0 0. 1 0. 3 0. 5 0. 8 0. 0 0. 1 0. 3 0. 5 0. 8 0. 0 0. 1 0. 3 0. 5 0. 8 – – – – – – – – – – – – – – – 0–2 2– 4 4–6 0–2 2–4 4–6 6–8 0–2 2–4 4–6 6–8 (d) T (e) F (f) K 0.848 0.848 0.848 0.68 0.68 0.68 0.512 0.512 0.512 0.344 0.344 0.344 2 2 S 0.176 0.176 S 0.176 D 0.008 0.008 D 0.008 M – 0.16 –0.16 M –0.16 N N – 0.328 –0.328 –0.328 – – –

0.496 0.496 0.496 – 0.664 –0.664 –0.664 – 0.832 –0.832 –0.832 –1 –1 –1 1 1 1 8 8 8 – – – 08 76 44 12 48 08 76 44 12 48 08 76 44 12 48 0.16 0. 6 0.16 0. 6 0.16 0. 6 0.832 0.664 0.496 0.328 0. 0 0. 1 0. 3 0. 5 0. 8 0.832 0.664 0.496 0.328 0. 0 0. 1 0. 3 0. 5 0. 8 0.832 0.664 0.496 0.328 0. 0 0. 1 0. 3 0. 5 0. 8 – – – – – – – – – – NMD S1 NMD S1 NMD S1 0–2 2–4 4–6 0–2 2–4 4–6 2.8–3 3–3.2 3.2–3.4 3.4–3.6 3.6–3.8 Fig. 4. Distribution of average Ellenberg’s Indicator Values in the first two dimensions of the NMDS ordination space (a–f)

138 J. FOR. SCI., 60, 2014 (4): 133–142 AIV unweighte weighted errorunwe errorweighted AIV N -0.34649 -0.4887614 0.259145 0.343666 AIV R -0.49547 -0.364978395 0.332521 0.333974 AIV F -0.1084 -0.08190916 0.100042 0.116039 AIV K 0.096196 0.15365566 0.10848 0.215058 AIV T -0.28278 -0.2989466 0.137451 0.198254 AIV L 0.198852 0.65711855 0.171398 0.31318

AIV L the different approaches to AIV calculation. In the course of the model diagnostics, distributions of AIV T residuals occurred to be close to normal, but stan- AIV K dardized residuals showed correlations with fitted values. For this reason, the true confidence inter- AIV F vals are probably wider than the calculated ones, AIV R and the unweighted AIV L, unweighted AIV F and unweighted AIV N weighted AIV R slope values cannot be trusted to weighted be significant. –1.5 –1 –0.5 0 0.5 1

Fig. 5. Ninety-five percent confidence intervals of the change within unweighted and weighted average Ellen- DISCUSSION berg’s Indicator Values over the 1961–2013 period The increased floristic resemblance of the 2013 ent. Although the historical relevés collected in the relevés to that typical of the forest sites described lowest of the studied tiers (i.e. the fir-beech zone) as oligotrophic and acidic suggests that in the top tend to occur on the left side of the ordination ranges of the Moravian-Silesian Beskids region the space, whereas those from near-peak elevations acidification levels of the soils are currently higher (the beech-spruce zone) seem to be found on the than during the post-war industrialization period. opposite side, the latter are underrepresented and This conclusion is further supported by the ob- no reliable judgement can be made. served AIV R (unweighted only) and AIV N (both The distribution of the average Ellenberg’s Indi- weighted and unweighted) decreases, and is con- cator Values (Fig. 4) logically matches the distri- sistent with the results of studies done in some oth- bution of the typological units: high AIV L values er parts of the Carpathian Mountains (Šamonil, are found on the right-hand side of the ordination Vrška 2007; Novotný et al. 2008; Durak 2010, space and can be associated with high-mountain 2011; Badea et al. 2012). Various authors provided stands, whereas the AIV R and N values increase different explanations of this phenomenon. Given toward the cluster of the nutrient-rich sites. The the proximity of large industrial centres – Ostrava, three remaining AIVs show a weak gradient. Třinec and Katowice agglomeration – to the re- The positions of the historical relevés collected in search area, it can be assumed that in the case of the 1960s (circles) do not differ much from the rele- the Beskids Mts. the observed changes are related vés originating from the 1970s (triangles). When to airborne pollutants, whose influence has been the positions of the 2013 relevés (marked with persisting despite decreased emission levels. Fol- squares) are considered, a roughly uniform pattern lowing the reasoning of Oulehle et al. (2010), the of change reveals itself: during the studied period, local soils could be expected to have high buffering the plant composition of the plots has shifted to- capacity. However, similarly like in the study done wards that typical of oligotrophic forests growing by Durak (2010), the acidification has been taking on acidic, calcium-poor soils (edaphic category K; place in spite of the occurrence of the flysch bed- low AIV R) and stands that occur at high elevations rock in the studied region. (high AIV L) or on gleyed soils (edaphic category O). The pollution impact has likely been augmented Lower is the share of the species commonly found by the maintenance of Norway spruce plantations at nutrient-rich localities (categories J and B; high on large areas of the mountain range. The acidifying AIV N). effect of this species is well documented (Augusto For four of the six AIVs a significant value of re- et al. 2002). The authors warn against establishing gression slope was obtained regardless of whether Norway spruce monocultures at the sites of high unweighted or weighted values were used (Fig. 5). acidic deposition, where the soil buffering capac- The confidence interval of the weighted AIV L ity has been exhausted. Resulting stands are sensi- change over the 1961–2013 period (95% CI 0.34 tive to disturbances and often show signs of health to 0.97 units, 133 df, P < 0.001) is wide, but far damage (Novotný et al. 2008; Badea et al. 2012). from zero, whereas the unweighted AIV L (95% CI The emission rates are beyond the control of for- 0.03–0.37, 133 df, P = 0.002) has reverse character- est managers, but the soil quality can be improved istics. In the remaining dependent variables with by changing the forest tree composition. The re- both significant slope values (i.e. AIV T, R and N), sults of unreplicated measurements of soil prop- the confidence intervals do not differ much under erties under first-generation forests in the Beskids

J. FOR. SCI., 60, 2014 (4): 133–142 139 region suggest that replacing Norway spruce with (Viewegh 1999), which were omitted by the pres- broadleaved species can support soil recovery in ent study. Were the evidence to be accepted, the re- this mountain range (Novák et al. 2012). A simu- sult would contrast with the predictions of increas- lation of future soil conditions in the Krušné hory ing drought frequencies at the top elevations of the Mts. (the Czech-German border) brought similar Moravian-Silesian Beskids Mts. near the end of the conclusions (Oulehle et al. 2007). The conversion 21st century (Hlásny et al. 2011). of Western Carpathian conifer forests into mixed ones between 1987 and 2005 was documented using remote sensing methods (Main-Knorn et al. CONCLUSIONS 2009), yet Norway spruce is still overrepresented in this region. In the previous decade it made al- In comparison with the period of the post-war most 75% of the Moravian-Silesian Beskids forest industrialization, the current composition of the cover (Vacek 2003). The conversion needs to be understorey plants in the top forest vegetation continued. zones of the Moravian-Silesian Beskids Mts. shows In contrast with soil acidification, changes in the an increased proportion of acidophilous and oli- light conditions of Carpathian forests have seldom gotrophic species. This finding suggests persisting been reported (Šamonil, Vrška 2007; Durak acidification of forest soils – probably in the re- 2010; Šebesta et al. 2011), and the observations do sult of past industrial emissions as well as forestry not agree with each other. In the present study, an policy promoting Norway spruce monocultures. A increased share of understorey species with high reduction of the extent of spruce plantations in fa- light Ellenberg’s Indicator Values was detected, but vour of stands composed of broadleaved tree spe- only the change of weighted AIV L was judged to cies should be considered as a way to support soil be statistically significant. The evidence obtained recovery. in the typological part of the analysis is also weak No vegetation shifts that could be related to the due to the insufficient number of observations global climate change were found. Some evidence of from stands growing in extreme conditions. This increasing light intensity and soil humidity is pres- kind of forests tends to have a status of small-scale ent, but it is weak due to an insufficient amount of specially protected areas and could not be covered data collected in wet and sparsely forested stands. by the present study. A shift of relevés in the ordi- In order to facilitate the monitoring of forest abiot- nation space toward the area occupied by observa- ic environment in the future, the establishment of tions from such stands was reported by Viewegh additional permanent plots – located at high eleva- (1999), who had explored vegetation changes of the tions, within small-scale specially protected areas same mountain range. Unfortunately, his analysis and on wet soils – should be considered. was based on a DCA ordination and the mentioned pattern occurred along the second axis, which tends to be deformed under this method. Acknowledgements Due to the small number of relevés collected at high-mountain stands the hypothesis about cli- The Forest Management Institute in Brandýs nad mate change-induced upslope shifts of plant popu- Labem offered the access to their network of phyto- lations could not be confirmed. On the contrary, sociological plots and provided the historical rele- the direction of the change of AIV T suggests slight vés. State Forests of the Czech Republic and the ad- cooling of the forest climate. The influence of the ministration of the Beskydy Protected Landscape global climate change on the abiotic conditions of Area permitted the data collection at the localities the forests in the studied area might have been too under their management. small to exceed the influence of other factors in the shaping of vegetation patterns. The translocation of the 2013 relevés towards References the part of the ordination space representing plant communities growing on gleyed soils suggests in- Augusto L., Ranger J., Binkley D., Rothe A. (2002): creasing water availability. Again, the available Impact of several common tree species of European tem- evidence is weak due to the insufficient number perate forests on soil fertility. Annals of Forest Science, of observations collected in wet forest stands. In 59: 233–253. the Moravian-Silesian Beskids Mts., habitats of Badea O., Bytnerowicz A., Silaghi D., Neagu S., Barbu I., this kind occur mainly in lower vegetation zones Iacoban C., Iacob C., Guiman G., Preda E., Se-

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Received for publication January 14, 2014 Accepted after corrections March 17, 2014

Corresponding author: Ing. Wiktor Żelazny, Mendel University in Brno, Faculty of Forestry and Wood Technology, Department of Forest Protection and Wildlife Management, Zemědělská 3, 61300 Brno, Czech Republic; e-mail: [email protected]

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