Download Herb layer dynamics of primeval fir

Ujházy, K.; Križová, E.; Vančo, M.; Freňáková, E.; Ondruš, M., 2005: Herb layer dynamics of primeval fir-beech forests
in central Slovakia. In: Commarmot, B.; Hamor, F. D. (eds): Natural Forests in the Temperate Zone of Europe – Values
and Utilisation. Conference 13-17 October 2003, Mukachevo, Ukraine. Proceedings. Birmensdorf, Swiss Federal Research Institute WSL; Rakhiv, Carpathian Biosphere Reserve. 193–202.
Herb layer dynamics of primeval fir-beech forests in central
Slovakia
Karol Ujházy, Eva Križová, Michal Vančo, Elena Freňáková and Miroslav Ondruš
Department of Phytology, Faculty of Forestry, Technical University in Zvolen, T. G. Masaryka 24, 960 53 Zvolen, Slovakia; [email protected], [email protected]
Abstract
Badínsky prales and Dobročský prales primeval forests are the oldest nature reserves in Slovakia, established
in 1913. Our current research in these primeval forests is aimed at the study of herb species dynamics during
the cyclic changes of fir-beech stands. We established 28 plots in different development stages of three
communities, sampled frequency data of the herb layer and measured the structure of the tree layer. Indirect
gradient analysis (DCA) showed separated clusters corresponding to different syntaxonomical units. Thus,
herb species dynamics during cyclic changes were community-specific, reflecting differences between habitats. We found relatively low species turnover during particular cycle in comparison to commercial forests.
Development stages were differentiated more by changing abundance or dominance of herb species. Generally, frequency values of most of herb species (closely correlated with the total herb cover) increased similarly as the number of trees and their estimated cover in the lowest layer (0–130 cm) – the traits representing
natural regeneration. On the other hand, they decreased with increasing sum of tree heights, as well as with
the number of trees and the cover of the main and/or understorey tree layer. However, in several herb species,
relationships deviating from this general rule were found. Especially the density of understorey trees seems
to be an important factor, limiting herb layer development more than shading of main storey trees.
Keywords: primeval forests, Eu-Fagenion, herb layer, cyclic changes, development stages, nature reserve,
Western Carpathians
1
Introduction
The Badínsky prales and Dobročský prales primeval forests belong to the oldest nature reserves in Slovakia.
At the time of their foundation in 1913 they were considered to be the best preserved sites of primeval firbeech forests in Slovakia. Since then, the ecosystems of these two reserves have been strictly protected and
maintained without direct human impact. Both reserves now offer good opportunity to study spatial and temporal dynamics of fir-beech forest ecosystems in Western Carpathians.
Changes of vertical and horizontal structure of a local tree stands have been already described very well.
The first research plots were founded in Dobročský prales in 1948 (Leibundgut 1959) to study structure,
development stages, production abilities and natural regeneration. Silvicultural research was conduced by
Kusák (1960), Priesol (1966), Korpeľ (1967, 1989) and Saniga (1999a). Structural characteristics and
changes in Badínsky prales were studied on permanent plots also by Korpeľ et al. (1991) and Saniga (1999b).
Results from these localities were used within well known descriptions of primeval forests life cycles and
development stages of Leibundgut (1978) and Korpeľ (1995).
On the other hand, the dynamics of whole phytocoenoses or whole ecosystems has not been described
and explained sufficiently till now. Randušková (1964) started to study changes of forest types of Dobročský
prales in course of stand development on parallel plots. She compared occurrence of typological units according to six development phases (originally distinguished by Leibundgut) and found different phytocoenose
- 193 -
types combinations in different development phases. She also noted surprisingly high abundance of spruce in
some parts of the reserve, which was not consistent with the herb layer and rejuvenation species composition.
Her results were published together with dendrometrical characteristics by Priesol and Randuška (1967).
Unfortunately, the research was not continued and, at present, it is not possible to identify the plots.
The main aim of our work was to contribute to the knowledge of herb layer development during the life
cycle of West Carpathian fir-beech forests. We want to describe the changes of species composition within
development stages, find the relations between herb layer changes and tree stand dynamics and to compare
development cycles of different community types.
We intended to describe a variability of plant community within forest type as a unit of potential natural
vegetation, following ideas of A. Zlatník, who defined a forest type as a complex of natural developmental
biocoenose stages as well as changed stages within the same permanent site conditions (Zlatník 1976). He
described system of Western Carpathian groups of forest types according to species composition of optimum
natural stand stages. He cannot describe all development stages of numerous forest types, but he emphasized
that the complete knowledge of particular types is possible only by means of long term observations of a
representative network of permanent plots (Zlatník 1961). Although numerous plots were established in Slovak beech forests, their temporal herb layer variability has not been studied sufficiently; species composition
of development stages of particular forest types has not been described yet. The lack of information on this
variability still causes problems in practical applications of forest typology in Slovakia.
2
Material and methods
Dobročský prales (48º 41´ N, 19º 40´ E) and Badínsky prales (48º 41´ N, 19º 03´ E) are located in central
Slovakia, central Western Carpathians. Both nature reserves are occupied mainly by fir-beech forests on
cambisol soil types. Badínsky prales (30.03 ha) is situated in Slovenské Stredohorie Mts. in the 4th beech
altitudinal vegetation zone (according to Zlatník 1959 classification), at 700–860 m a. s. l., on the volcanic
andesite bedrock, with optimal conditions for beech (Fagus sylvatica) which reaches maximal height of
about 45 m.
Dobročský prales (49.9 ha of core area, 103.9 ha whole area of reserve) in Veporské vrchy Mts. is on
granite bedrock, climatically shifted to the 5th fir-beech altitudinal vegetation zone in 720–1000 m a. s. l.,
with a natural occurrence of spruce (Picea abies) and optimal conditions for fir (Abies alba). Fir reaches the
height of 58 m and above ground biomass of 51.3 m3 (Randuška, Priesol 1967).
According to the Slovak typological classification system (Hančinský 1972), fir-beech forest communities of Badínsky prales belong mainly to groups of forest types Fagetum typicum and Fagetum pauper (Križová 2000), whereas communities of Dobročský prales are classified as Abieto-Fagetum and partly as
Fageto-Aceretum. According to Zürich-Montpellier school, herb-rich communities of Badínsky prales are
close to Asperulo-Fagetum Sougnez et Thill 1959 association, nudal types are classified as Dentario bulbiferae-Fagetum Zlatník 1935. Both communities are similarly formed by beech stands with regular admixture
of fir. Herb-rich fir-beech stands of Dobročský prales belong to the Dentario enneaphylli-Fagetum salvietosum glutinosae Oberdorfer ex W. et . A. Matuszkiewicz ex Guzikova et Kornaś 1969. They are dominated by
beech and fir, with a regular admixture of sycamore (Acer pseudoplatanus), rarely also with spruce (Picea
abies), elm (Ulmus glabra), ash (Fraxinus excelsior) and maple (Acer platanoides).
In each reserve, 14 square or rectangular plots, 400 m2 in size have been established (in Badínsky prales
in 2001 and in Dobročský prales in 2002). Plots were placed with the aim to cover different stages of tree
stand development after Korpeľ et al. (1991). Series of plots were set up in surroundings of optimum stages
(with well-developed herb layer) with similar site conditions. Number of plots in one series (from three to
ten) and number of series were limited by heterogeneity of site conditions, size of reserves and, mainly in
Badínsky prales, by lack of optimum
stages. Overall we have established one
series with three plots in Fagetum pauper (Fp), three series with eleven plots
in Fagetum typicum (Ft) and series 14
plots in Abieto-Fagetum (AF).
Five additional relevés from notfixed plot (nr. 17–21) from Badínsky
prales and two fixed plots (nr. 3, 4) from
Figure 1. Sample plots with 200 squares on belt transects.
- 194 -
buffer zone of this reserve were used for some analyses.
The plots were stabilised in corners by wooden poles. Following data were recorded on each plot:
- abundance and dominance of all vascular plant species using Zlatník´s estimating scale (Zlatník 1976)
- vertical structure by estimating woody species cover in each of seven layers: 1 – over-storey; 2 – main
storey; 3 – under-storey (higher than 1/2 of main storey); 4 – from 1.3 m to 1/2 of main storey; 51a – from
0.2 to 1.3 m; 51b < 0.2 m; 52 – juvenile trees
- frequencies of herb layer species (including woody species) recorded on two-hundred 0.25 m2 square
subplots located along systematically placed belt transects within the plot (Fig. 1)
- absolute abundance of tree species (separately according to vertical layers) counted on each transect
- diameter and height of all trees with diameter at breast height (dbh) > 7 cm
The classification of development stages follows Korpeľ (et al. 1991) with English equivalents (growing,
optimum, decaying stage) according to Neuert (et al. 2001). Names of typological units are used according to
Hančinský (1972). Plant species nomenclature follows Marhold, Hindák (1998). Rubus hirtus was determined as a broad species R. hirtus Waldst. et Kit. (Dostál, Červenka 1991). Species of Dryopteris carthusiana agg. were not determined, however, in most cases it should be Dryopteris dilatata.
The relevés were stored and processed using the TURBOVEG programme package (Hennekens 1996a)
and MEGATAB programme (Hennekens 1996b). Similarity of all relevés (from both localities) was analysed
by TWINSPAN divisive classification method (Hill 1979). Further analyses were based mainly on herb species frequencies. CANOCO programme (ter Braak & Šmilauer 1998) was used for indirect gradient analysis
(Detrended Correspondence Analysis – DCA) and for testing the tree stand characteristics in relation with
herb layer variability by Monte Carlo permutation test. Statistica 6.0 software was used for correlation analysis using Spearman rank correlation coefficient.
3
Results
Samples were taken in three vegetation units with quite similar species composition of herb layer. 28 out of
57 herb species were found in both reserves. Only 19 of these common species occurred at more than 50 %
of plots in one reserve (Actaea spicata, Athyrium filix-femina, Dentaria bulbifera, Dryopteris dilatata,
Dryopteris filix-mas, Galeobdolon luteum, Galium odoratum, Geranium robertianum, Impatiens nolitangere, Mercurialis perenis, Oxalis acetosella, Paris quadrifolia, Prenanthes purpurea, Rubus idaeus, Salvia glutinosa, Sanicula europaea, Senecio ovatus, Urtica dioica, Viola reichenbachiana). All dominants (in a
single plot) belong to the most frequent species. Differentiating species between the two reserves were generally both less frequent and dominant, with the exception of Rubus hirtus in Badínsky prales, Dentaria enneaphyllos, Gymnocarpium dryopteris and Stellaria nemorum in Dobročský prales.
The first division of TWINSPAN classification strictly separated the two localities (Fig. 2), which is
clearly the consequence of the numerous differentiating species and down-weighting of cover values of the
relevé nr.:
locality:
herb layer cover (%):
2
3
4
7
85
65
25
92
12
13
14
1
8
11
Dobročský prales
85 68 38 15
35
5
9
6
10
12
28
12
10
development stage: dec opt dec dec dec opt gro gro gro
gro
opt
opt
gro opt
typological unit:
AF
AF
AF
AF AF* AF* AF AF* AF
AF*
AF
AF
AF
AF
vegetation stage:
trophic series:
5
B
5
B
5
5
5
5
5
5
B B(/C)B(/C) B B(/C) B
5
B
17
20
21
11
16
12
7
10
80
45
20
6
4
8
15
14
2
9
13
18
19
1
3
5
Badínsky prales
35 2.5 2.5 3
2.5
5
20 10 0.5
4
30 25
9
dec opt opt dec opt opt "gro" dec opt gro gro opt dec
gro
gro gro
dec
gro opt "opt"
gro
Ftil FAc Ft Ft Ft Ft Fp Ft Ft Fp Ft Fp Fp
Ft
Ft
Ft Ft Fp
FrAc Ft
Ft
4
4
4
4
5
4
4
4
4
4
4
4
4
B
B/C B/C B
B
B
B
B
B
B
B
B
B
C
75
65
80
30
30
Figure 2. TWINSPAN classification of phytosociological relevés from all sample plots based on transformed
cover values of herb layer species. Abbreviations: opt = optimum, dec = decaying, gro = growing stage, B =
mesotrophic, B/C = heminitrophitic, C = nitrophytic.
- 195 -
most frequent and dominant species. The second division approximately separated herb rich stages and
stages with low herb layer cover (independent of the
community type). Three relevés from heminitrophilous
and nitrophilous communities of Badínsky prales
(Fagetum tiliosum, Fageto-Aceretum and FraxinetoAceretum) were allocated to the cluster of mesotrophic
communities (Fagetum typicum) with well-developed
herb layer. Development stages of Fagetum typicum
with low herb cover were, on the other hand, allocated
to the cluster of the nudum type (Fagetum pauper).
Herb-rich relevés of the decaying stages of mesotrophic Abieto-Fagetum type were clustered together with
relevés from heminitrophillous type (Abieto-Fagetum*
– Fig. 2, 3) in Dobročský prales.
The application of indirect gradient analysis (DCA)
to frequency data (Fig. 3; each plot was represented by
sampled species frequencies; rare species with presence on less than three plots were omitted) yielded
better results (from the point of view of vegetation
dynamics). Plots were ordered into clusters according
to the community type (vegetation unit). With exception of plot 11 from the Dobročský prales, plots of
different communities were not mixed together and
vegetation dynamics could thus be analysed for each
community separately.
Figure 3. Ordination of sample plots according to herb
species frequencies (DCA). B – plots from Badínsky, D
– Dobročský prales.
3.1 Herb species dynamics during Abieto-Fagetum development cycle
In this paper, we are focusing on the best documented series of Abieto-Fagetum from Dobročský prales. Plots
1–11D were considered.
Tabular synthesis (Table 1) shows that with exception of Galeopsis sp. occuring only in decaying stage,
there are no distinctive species for each stage. However, there seem to be few species which can be used for
negative floristic differentiation of the growing stage (Senecio germanicus, Galeopsis sp., Circaea alpina).
According to these results, species change their frequency but most of them are present during the whole
cycle. Increasing frequency values can be used for estimating the species optimum and each stage can be
characterised by species that reach their ecological optimum in this given stage. Two ferns – Gymnocarpium
dryopteris and Dryopteris dilatata – have optimum in growing stage. Two other species which can grow on
thick raw-humus layer or directly on dead wood (Oxalis acetosella, Dryopteris dilatata) are also frequent in
the growing stage. However, their highest frequency values were recorded in the decaying stage.
Viola reichenbachiana is the only species with a distinct optimum in the optimum stage. Another group
of species (Galium odoratum, Galeobdolon luteum, Stellaria nemorum, Salvia glutinosa) has high (the highest) frequencies in both the optimum and the decaying stage. The group of nitrophytic species has increasing
values through the whole cycle from the growing through the optimum to the decaying stage. The last group
of taxa with rare records in the growing and the optimum stage (Urtica dioica, Senecio sp., Rubus idaeus,
Galeopsis sp.) is characterised by tall-growing plants and is associated with humus decomposition in decaying stage.
DCA indirect gradient analysis (Fig. 4, 5) provided the differentiation of species to the three groups corresponding to three stages of development cycle. Sample plots as well as species are arranged to the circular
pattern according to the first two axes of ordinal space. This spatial pattern should have also temporal character, if we expect that position of sample plots will rotate counter-clockwise in the future.
- 196 -
Table 1. Tabular species frequencies differentiation of Abieto-Fagetum development stages. Frequencies
higher than average are bolded. Less frequent species without distinct dynamic pattern were not included
(Asarum europaeum, Dentaria eneaphylos, Paris quadrifolia, Polygonatum verticillatum, Veronica montana,
Chrysosplenium alternifolium, Stachys sylvatica).
6d
d e ve l o p m e n t s t a g e
h e r b l a y e r c o ve r ( % )
c a n o p y (% )
t r e e l a y e r c o ve r ( % ) - 1 , 2
- 3, 4
- 5
G y m n o c a rp iu m d ry o p t e ris
D ry o p t e ris c a rt h u s ia n a a g g .
O x a lis a c e t o s e lla
A t h y riu m f ilix -f e m in a
D e n t a ria b u lb if e ra
D ry o p t e ris f ilix -m a s
A n e m o n e n e m o ro s a
P re n a n t h e s p u rp u re a
V io la re ic h e n b a c h ia n a
M y c e lis m u ra lis
S a n ic u la e u ro p a e a
G a liu m o d o ra t u m
G a le o b d o lo n lu t e u m
S t e lla ria n e m o ru m
S a lv ia g lu t in o s a
A c ta e a s p ic a ta
G e ra n iu m ro b e rt ia n u m
I m p a t ie n s n o li-t a n g e re
M e rc u ria lis p e re n n is
R u b u s id a e u s
U rt ic a d io ic a
S e n e c io o v a t u s
S e n e c io g e rm a n ic u s
C irc a e a a lp in a
G a le o p s is s p .
12
55
4
86
4
4
18
19
4
5
2
2
1
8
1
1
1
0
11d
5d
g ro w in g
15
28
95
65
8
65
98
53
3 ,5
6
10
51
12
2
1
1
10
6
34
20
10
4
2
1
7
42
1
2
2
7
11
25
2
3
4
5
1
2
4d
9d
25
35
31
54
5
28
30
25
41
8
5
28
23
18
5
8
2
2
2
1
27
24
2
1
1
6
7
6
8
2
1
1d
8d
o p tim u m
38
35
90
95
80
85
34
35
4
5
20
6
73
16
9
6
1
3
33
11
10
5
3
44
14
18
5
12
12
4
5
30
2
1
2
4
1
1
84
51
29
3
13
59
57
14
4
1
4
9
2
3
1
1
1
1
8
1
4
9
9
2
1
1
7d
2d
d e c a y in g
92
85
40
25
20
45
12
42
5
14
3d
4
3
85
20
9
5
1
12
58
23
20
14
9
3
1
2
7
1
2
82
69
6
7
2
72
94
44
35
3
1
3
62
51
18
6
2
1
54
68
6
3
2
68
39
13
48
16
2
1
7
2
65
55
25
37
24
1
65
56
4
7
2
24
54
24
26
5
3
1
2
5
Figure 5. DCA differentiation of species according to stages:
triangle = growing, circle = optimum, cross = decaying.
Figure 4. DCA ordination of sample plots following
circular character of Abieto-Fagetum development.
- 197 -
Oxa ace
Dry fil
Ste nem
Act spi
Asa eur
San eur
Vio rei
Sal glu
Gal lut
Ger rob
Imp nol
Gal odo
Sen ova
Urt dio
Mer per
Den bul
3.2 Relationships between herb species frequencies and tree layers traits
We used Monte Carlo permutation test to study changes of herb species frequencies according to several
measured or estimated stand variables. The variables that best explained the variance of herb species frequencies are: abundance in 51a tree layer – 10.1 % (of total variance explained); cover of 3 + 4th tree layer –
9.7 %; abundance of beech in 51a – 8.9 %; sum of tree heights – 7.4 %; abundance in 51 tree layer – 7.3 %;
cover of 4th layer – 7.2 %.
Partial correlations of individual species with tree
Table 2. Significant (bold) correlations of optimum
layer traits using Spearman rank correlation coeffistage species frequencies with tree layers traits and
cient allowed us to distinguish two groups of herb
total herb layer cover.
species with different responses to the traits (Table 2
and 3).
Optimum stage species seems to be adapted to medium-shady conditions in “hall” mature stands with a
total herbs cov er
0.1 0.8 0.6 0.4 0.8 0.7
tree mainstorey cover
0.6 0.6 0.6 0.5 0.5 0.4
closed canopy. Dense understorey (3+4) characteristic
mainstorey abundancy
0.5 0.5 0.7 0.6 0.4 0.4
for the growing stage probably suppressed some herb
understorey abundancy -0.6 -0.6 -0.1 -0.2 -0.4 -0.6
species, especially Dentaria bulbifera and Galium
juvenile trees abundancy -0.4 -0.3 -0.3 -0.3 -0.6 -0.8
odoratum.
On the contrary, decaying stage species
Table 3. Significant (bold) correlations of decaying and growing
such as Urtica dioica, Impatiens nolistage species with tree layers traits, total herb layer cover and sum
tangere or Geranium robertianum (conof all species frequencies on plot.
nected with a higher light and subsequently
also nutrient supply) were recorded with the
highest frequencies when canopy was open
(low cover and abundance of mainstorey and
total herbs cover
0.8 0.7 0.4 0.9 0.9 1.0 0.9 0.7 0.5 0.8
5 1 cover
0.6 0.8 0.8 0.6 0.7 0.5 0.9 0.7 0.6 0.6
understorey trees negatively indicated by
5 1a cover
0.6 0.9 0.7 0.6 0.7 0.5 0.7 0.7 0.6 0.6
decreasing sum of trees heights of 1–3 lay5 1b cover
0.6 0.7 0.8 0.5 0.5 0.4 0.8 0.4 0.4 0.4
ers) and regeneration was in progress (posi5 1a abundancy
0.5 0.3 0.2 0.7 0.8 0.6 0.7 0.1 0.3 0.5
tive correlations with abundance and the
sum of frequencies 0.4 0.2 0.0 0.6 0.6 0.6 0.2 0.1 0.2 0.5
cover of trees < 1.3 m).
sum of tree heights -0.4 -0.7 -0.8 -0.6 -0.7 -0.5 -0.9 -0.5 -0.4 -0.4
Dryopteris
filix-mas
and
Oxalis
acetosella, the two of “growing stage species” have similar “correlation pattern” as decaying species. Although they have higher frequencies in growing stage than other species, their frequency values are highest in
decaying stage, too.
frequency (%)
3.3 Interspecific correlations
Interspecific relations could play an important role, at least in the later optimum and in the decaying stage
with high herb layer cover, since competition might gain importance in these stages.
We found a group of strongly positively correlated species (Geranium robertianum, Galeobdolon luteum,
Impatiens noli-tangere, Mercurialis perennis, Galium odoratum; Table 4). However, correlation of their
frequencies can most likely be explained by similar reactions to changing ecological conditions.
Galium odoratum and Galeobdolon luteum
100
(the
second and third most frequent species of
90
Geranium robertianum
80
local
beech forest with Oxalis acetosella being the
Impatiens noli-tangere
70
most
frequent) are typical for optimum stage.
Salvia glutinosa
60
R = 0,72
They are grouped with three nitrophytic species
50
40
R = 0,62
whose frequency increases more rapidly in the
30
R = 0,72
decaying stage. Differences in the development of
20
10
frequencies of these closely correlated species to
0
changing light and nutrient levels from the grow0
20
40
60
80
100
ing to the decaying stage are expressed by the
frequency of Galium odoratum (%)
regression on Figure 6.
The only significant negative correlation
Figure 6. Exponential regression of frequencies of
(weak
negative power regression) was observed
Galium odoratum and three nitrophytic species.
between species Dryopteris dilatata and Stellaria
nemorum, but Stellaria nemorum is not a very frequent species of those communities.
2
2
2
- 198 -
Table 4. Matrix of Spearman rank correlations of herb species frequencies from all plots of Dobročský prales. Significant
correlations (on p<0.05) are indicated in bold.
Geranium robertianum
Galeobdolon luteum
Impatiens noli-tangere
Mercurialis perennis
Galium odoratum
Salvia glutinosa
Oxalis acetosella
Stellaria nemorum
Dryopteris filix-mas
Rubus idaeus
Athyrium filix-femina
Viola reichenbachiana
Sanicula europaea
Dentaria bulbifera
Dryopteris dilatata
4
Ger rob Gal lut Imp nol Mer per Gal odo Sal glu Oxa aceSte nem Dry fil Rub ida Ath fil Vio rei San eur Den bul Dry dil
1.0
0.8
0.9
0.8
0.8
0.6
0.6
0.5
0.5
0.3
0.4
0.4
0.3
0.2
0.1
0.8
1.0
0.8
0.7
0.7
0.6
0.7
0.6
0.3
0.4
0.3
0.5
0.3
-0.1
0.0
0.9
0.8
1.0
0.9
0.8
0.6
0.7
0.3
0.3
0.4
0.2
0.3
0.1
0.3
0.0
0.8
0.7
0.9
1.0
0.7
0.4
0.6
0.4
0.5
0.2
0.0
-0.1
-0.1
0.3
0.0
0.8
0.7
0.8
0.7
1.0
0.9
0.4
0.6
0.3
0.1
-0.1
0.5
0.4
0.2
-0.2
0.6
0.6
0.6
0.4
0.9
1.0
0.3
0.5
-0.1
0.0
-0.1
0.6
0.5
-0.1
-0.1
0.6
0.7
0.7
0.6
0.4
0.3
1.0
0.0
0.3
0.2
0.4
0.4
0.4
-0.2
0.4
0.5
0.6
0.3
0.4
0.6
0.5
0.0
1.0
0.3
0.1
0.0
0.2
0.1
0.4
-0.7
0.5
0.3
0.3
0.5
0.3
-0.1
0.3
0.3
1.0
0.0
0.4
0.0
-0.2
0.4
0.0
0.3
0.4
0.4
0.2
0.1
0.0
0.2
0.1
0.0
1.0
0.8
-0.4
-0.6
-0.1
-0.1
0.4
0.3
0.2
0.0
-0.1
-0.1
0.4
0.0
0.4
0.8
1.0
-0.2
-0.2
-0.1
0.5
0.4
0.5
0.3
-0.1
0.5
0.6
0.4
0.2
0.0
-0.4
-0.2
1.0
0.9
0.0
-0.2
-0.6
-0.2
0.9
1.0
-0.2
0.0
0.3
0.3
0.1
-0.1
0.4
0.5
0.4
0.1
-0.2
0.2
-0.1
0.3
0.3
0.2
-0.1
-0.2
0.4
0.4
-0.1
-0.1
0.0
-0.2
1.0
-0.4
-0.1
0.4
-0.7
0.0
-0.1
0.5
-0.2
0.0
-0.4
1.0
0.1
0.0
0.0
0.0
-0.2
Discussion
The processes studied in both primeval forests can be considered as fluctuations in a broad sense according to
Miles (1979), or as an internal community dynamics (van der Maarel 1996), when vegetation keeps the same
overall appearance. Changes in herb layer are influenced by beech and fir population dynamics, their mortality and regeneration. Whole process can be interpreted as gap or patch dynamics (depending on size of canopy gaps).
Especially in Dobročský prales we met small-scale horizontal pattern of development stages given by the
death of one or couple of old canopy trees. Regeneration of beech is quite rapid in this small openings with
the width most frequently not more than 20 m. This characteristic feature can explain surprisingly low species turnover during the development cycle – we noted only five taxa to grow only in decaying stage (Chrysosplenium alternifolium, Galeopsis sp., Festuca altissima, Fragaria vesca, Pulmonaria obscura – the latter
three were recorded only within complete relevés; all of them were rare species). Seven of decaying stage
species were missing on growing stage. Lot of other so called “clearcut flora” species were not found on any
plot, however, we tried to find the most opened stages.
FAJMONOVÁ (1986), who studied clearcut plant communities on sites of fir-beech forests on crystalic
bedrock of four mountain ranges within inner Western Carpathians, recorded these diagnostic species of
Epilobietea angustifolii class: Rubus idaeus, Epilobium angustifolium, Senecio germanicus (the highest constancy), Senecio ovatus, Omalotheca sylvatica, Fragaria vesca, Eupatorium cannabinum (lower constancy).
RANDUŠKA et al. (1985) and KRIŽOVÁ (1991) compared herb layer species composition of mature stands and
clearcuts of the Abieto-Fagetum group of forest types on 9 pairs of plots in the Nízke Tatry and Kremnické
vrchy Mts. (Central Slovakia). On clearcuts they found 11–43 species in addition to mature stands with
closed canopy. Among them, there were some ruderal species (as Tussilago farfara, Cirsium spec. div.),
several heliophilous species of oak forests (Hypericum perforatum, Steris viscaria) and a number of a typical
clearcut species not recorded in the reserve (Chamaerion angustifolium, Digitalis grandiflora, Eupatorium
cannabinum, Fragaria vesca). Chamaerion angustifolium and Rubus idaeus were always present on clearings of beech forests in Romanian Carpathians (RESMERITA 1982). Rubus idaeus is the most characteristic
species in the gaps of the Dobročský prales reserve, but Chamaerion angustifolium is missing. Also Senecio
nemorensis agg. does not play important role here in comparison with well-known situation in clearcuts or
opened canopy stands of managed fir-beech forests. According to RANDUŠKA et al. (1985) and KRIŽOVÁ
(1991), it belongs among the most frequent dominants of clearcuts (along with Rubus idaeus, Urtica dioica,
Galium odoratum, Stachys sylvatica, Oxalis acetosella, Geranium robertianum and Rubus hirtus).
Unfortunately, there are no comparable works dealing with herb layer changes within development cycle
of primeval forests communities from Western Carpathians. From the Central European region, we can compare our results with long-term studies from some forest reserves of Czech Republic. VRŠKA et al. (2002)
studied changes of Dentario enneaphyli-Fagetum after 21 years on exactly recovered permanent plots. On
most of plots of comparable community type (6D1) in the Žákova hora Mt. reserve (Českomoravská
vrchovina uplands), they observed decreased total herb cover due to the developing complex tree stand structure and hence a greater shade. Decreased cover degree showed Rubus idaeus, Urtica dioica, Mercurialis
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perennis, Senecio nemorensis, Galium odoratum, Galeobdolon luteum, Oxalis acetosella and Viola reichenbachiana. Only Dentaria bulbifera and Dryopteris carthusiana had slightly increased covers.
These results are consistent with our findings, when the above mentioned “increasing” species belong to
growing stage species (Fig. 5, Table 1) tolerating a high tree density and the “decreasing” species we classified as optimum and decaying stage species negatively correlating with the sum of tree heights – the trait
reflecting mainly a high number of understorey trees (Table 3). Mercurialis perennis, Galium odoratum,
Galeobdolon luteum and Oxalis acetosella also belong to the group of species with the most close reaction to
the stand changes (Table 4) in the Dobročský prales.
VRŠKA et al. (2002) explain the changes mainly by changing light conditions. Probably this is the most
important factor, however, our results show also two other important features. Negative reaction of most of
species to increasing beech stand density can be explained also by increasing root competition. SLAVÍKOVÁ
(1986) presented an example, when cut across of beech roots enabled development of herb layer under the
same light conditions of Fagetum nudum community. Other feature is live strategy and life form. The most
rapid development showed polychoric and autochoric terophytes Geranium robertianum and Impatiens nolitangere (cf. JURKO 1990), followed by species with vegetative reproduction ability (Galium odoratum,
Galeobdolon luteum and others). On the other hand, long living perennial ferns survived without a big
change of abundance through the whole cycle (Table 1). RESMERITA (1982) ranked Athyrium filix-femina,
Dryopteris filix-mas and Dryopteris carthusiana to the same group of shade tolerant species which can also
survive long time on completely opened clearcuts.
We were interested also on whole community changes and variability. Although species turnover is relatively low, the composition undergoes considerable changes during the development cycle of the studied
communities. The comparison of three different cycles by means of frequency values ordination (Fig. 3)
shows that these changes do not exceed community type. Within this cyclic process, ordination seems to be a
more efficient method to classify the stages than classification methods such as TWINSPAN (Fig. 2), whose
results reflects more floristic differences than developmental trends.
This result is important for the application in syntaxonomic classification systems used in Slovakia,
which do not consider variability between development stages enough. We can see (Table 1), that communities of development stages are differentiated more by dominance os species than by other species occurence.
Thus, especially when dominance is considered to be important classification criterion, some development
stages of the same community can be classified separately or together with stages of the other community
(Fig. 2)
Units of the Slovak typological school are differentiated both by the presence and the cover of diagnostic
species. Thus, at the same site of Abieto-Fagetum, the floristic composition can vary in the range of two or
three groups of forest types. In particular, herb layer with the dominance of nitrophytic species, such as Geranium robertianum and Impatiens noli-tangere (cf. Križová, Nič 1997), in the decaying stage of AbietoFagetum would be improperly classified as Fageto-Aceretum.
Even if we speak about a broad association as Dentario enneaphylli-Fagetum from Fagion alliance of the
floristic-based classification system of Zürich-Montpelliér school, its decaying stage in Dobročský prales
with increasing dominance of nitrophytes seems to be closer to the heminitrophilous Mercuriali-Fraxinetum
association (sensu Husová in Moravec et al. 2000) from Tilio-Acerion alliance.
Consequently, the incorrect estimation of development stage can lead to the improper classification results. In the case of forest typology, it can have negative consequences in forest management practice, such
as plantations of less suitable tree species.
5
Conclusions
Some common features of herb layer changes during natural fir-beech primeval forests development were
found:
- In both reserves studied in Central Slovakia, frequencies of most of the herb species had the tightest associations with the total herb cover (estimated on a whole plot), number of individual trees in the lowest
(5th) layer (0–130 cm) and estimated 5th layer tree cover (whole plot). The decline of frequencies was explained by the increasing values of summed tree heights (dbh > 7 cm), a number of individuals and the
cover of particular tree layers (mainstorey and understorey).
- Temporal development of plant populations as well as their relationships with tree layer traits are nonlinear; we can find groups of species with similar reaction to the changes of tree stand.
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- There is a relatively high frequency of ferns (Dryopteris carthusiana agg., D. filix-mas, Athyrium filixfemina) and Dentaria bulbifera in dense and shady growing stages and rapid frequency increase of nitrophytic (Geranium robertianum) or so called “clear cut species” (Rubus idaeus) in opened gaps of decaying stage.
- There is a low species turnover during the development cycle of fir-beech primeval forest in comparison
to managed forests with clear-cut stages.
- The highest vascular plant diversity of fir-beech primeval forests is in the beginning of decay stage and in
the phase of beech regeneration.
However, behaviour of particular species and trees/herbs relations differ between the two localities and
community types as well. In Badínsky prales, the 4th layer (from 1.3 m to half of main storey height) featured
the strongest negative influence on the herb layer species frequencies, whereas in Dobročský prales, it was
the 3th layer (understorey).
In best described series of Abieto-Fagetum in Dobročský prales very clear gradient from oligotrophic,
mesotrophic to eutrophic and nitrophytic species between the growing and decaying stage was observed.
Similar gradient was not so obvious in different conditions of Badínsky prales.
Dentaria bulbifera was not significantly positively related to any other species but in Badínsky prales it
showed significant (p < 0.05) negative correlation with some of them (Impatiens noli-tangere, Galium odoratum and Dryopteris filix-mas). We distinguished groups of species with similar reactions to tree stand
structure changes and with tight positive correlations between their frequency values as well, but they partly
differ between two localities (Galeobdolon luteum, Geranium robertianum, Impatiens noli-tangere and Galium odoratum in Dobročský prales; Galium odoratum, Geranium robertianum and Oxalis acetosella in
Badínsky prales).
In conclusion, we scan say that different herb species dynamics were related to different community
types, reflecting differences in habitats and available species pool of particular phytogeographical region, as
shown by indirect gradient analysis of the same herb frequencies data set. DCA results showed separated
clusters of the above-mentioned community types. Ordering of plots according to the development dynamics
could be recognized only within particular clusters. The variability of species composition within the development cycle does not seem to exceed limits of community type, although it markedly exceeds “static” perception of community represented only by single phytosociological relevé of mature forest stand.
6
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Acknowledgements
This paper was supported by the Slovak Grant Agency for Science (grant No. 1/0629/03).
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