HACQUETIA 3/1 • 2004, 61–73


THE BEECH FOREST VEGETATION OF THE 
CEROVÁ VRCHOVINA MTS. (SOUTHERN SLOVAKIA) 

Karol UJHÁZY*, Richard HRIVNÁK**, Eva BELANOVÁ***
, 
Blažena BENČAŤOVÁ*


Izvleček 

Leta 2001 smo izvedli terenske fitosociološke raziskave sestojev, na gori vulkanskega izvora Cerová vrchovina 
v katerih dominira vrsta Fagus sylvatica. Uporabili smo standardno srednjeevropsko fitocenološko metodo. Za 
ločevanje posameznih združb smo naredili tabelarično sintezo s pomočjo numerične ločitvene klasifikacije 
(TWINSPAN) in indirektne gradientne analize (PCA). Znotraj zveze Fagion smo ločili štiri asociacije: Carici 
pilosae-Fagetum, Dentario bulbiferae-Fagetum, Melico-Fagetum in Asperulo-Fagetum. V zvezo Tilio-Acerion smo uvrstili 
štiri rastlinske sintaksone: Roso pendulinae-Tilietum cordatae, združba Mercurialis perennis-Fagus sylvatica, združba 
Athyrium filix-femina-Fagus sylvatica in združba Dryopteris filix-mas-Fagus sylvatica. Delno oligotrofne združbe smo 
uvrstili v asociacijo Luzulo-Fagetum in zvezo Luzulo-Fagion. Obravnavali smo vpliv številnih ekoloških faktorjev 
na vrstno sestavo. Relief, kamnitost in vsebnost humusa so med najpomembnejšimi. 

Abstract 

Field phytosociological research of the Fagus sylvatica dominated forests in the volcanic Cerová vrchovina 
Mts. was carried out in 2001. Standard methods of the Zürich-Montpellier approach were applied. Tabular 
synthesis with numerical divisive classification (TWINSPAN) and indirect gradient analysis (PCA) were used 
to differentiate particular communities. Within the Fagion alliance, four associations were recognized: Carici 
pilosae-Fagetum, Dentario bulbiferae-Fagetum, Melico-Fagetum and Asperulo-Fagetum. Four different plant communities of the Tilio-Acerion alliance were detected: Roso pendulinae-Tilietum cordatae, Mercurialis perennis-Fagus 
sylvatica community, Athyrium filix-femina-Fagus sylvatica community and Dryopteris filix-mas-Fagus sylvatica community. Semioligotrophic communities were classified as Luzulo-Fagetum within the Luzulo-Fagion alliance. 
Influences of several ecological factors on species composition are discussed. The type of relief, stoniness and 
humus content of topsoil seem to play the most important role. 

Ključne besede: Slovaška, vegetacija, Fagetalia, klasifikacija, gradientna analiza
Key words: Slovakia, vegetation, Fagetalia, classification, gradient analysis


1. INTRODUCTION on the northern slopes or shady valleys. On fully 
developed soils, communities of the Fagion alliance 
The Cerová vrchovina Mts. forms a volcanic moun-occur. On small patches of scree and boulder fields 
tain range situated in the southern part of the with a specific mesoclimate grow the communities 
Western Carpathians. Under the influence of a of the Tilio-Acerion alliance. 
relatively warm and dry climate, communities with The study area belongs to those regions of Slothe dominance of Fagus sylvatica are restricted only vakia which have been relatively well investigated 

* Department of Phytology, Faculty of Forestry, Technical University of Zvolen, Masarykova 24, SK-960 53 Zvolen, Slovakia, 
e-mail: (a) ujhazy@vsld.tuzvo.sk, (b) bbencat@vsld.tuzvo.sk 
** Institute of Botany, Slovak Academy of Sciences, Dúbravská cesta 14, SK-845 23 Bratislava, e-mail: richard.hrivnak@savba.sk 
*** State Nature Conservancy of the Slovak Republic, Protected Landscape Area Cerová vrchovina, Svätoplukova 40, SK-979 01 
Rimavská Sobota, e-mail: belanova@sopsr.sk 
61 


HACQUETIA 3/1 • 2004 

from the point of view of floristics. The basic reviews about flora of vascular plants of the Cerová 
vrchovina Mts. and surroundings were published 
by Hendrych (1959, 1963, 1968) and Holub & Moravec (1965). On the other hand, only a few phytosociological papers discussing local forest vegetation exist. The occurrence of the associations Carici 
acutiformis-Alnetum glutinosae Scamoni 1935 and Aegopodio-Alnetum glutinosae Šomšák 1961 was presented by Balázs (1996). Csiky & al. (2001) described 
a new forest community of block forest – Roso pendulinae-Tilietum cordatae – from this area. 

2. STUDY AREA 
The Cerová vrchovina Mts. is situated at the southern edge of central Slovakia, on the border with 
Hungary. In the northwest, it is bordered by the 
Ipel’ river and in the northeast by the Rimava river. 
From the south, the studied area is delimited by 
the Hungarian state boundary (Fig. 1). 


Figure 1: Location of studied area 
Slika 1: Lokacija raziskovanega območja 

The highest point of the Cerová vrchovina Mts. 
is Karanč (725 m a. s. l.). The lowest point of the Slovak part is near the Vlkyňa village at 155 m a. s. l. 

Steep conic volcanic hills are characteristic for 
this region, rising from the smooth relief of sedimentary rock. Volcanic hills and mountain ridges 
are built mainly by Pliocene-Pleistocene basalt lava 
f lows, agglomerates and lapilli tuffs. Other volcanic rocks, such as andesites (Karanč and Šiator hills) 
or rhyodacite tuffs and tuffites (situated on the western part) occur rarely as well. However, the largest 
part is covered by Miocene sediments, most frequently with disintegrating sandstones. 

From the phytogeographical point of view this 
Carpathian region belongs to the Matricum district 
within the Pannonicum phytogeographical region 
(Futák 1966, cf. Molnár 1999). 

The existence of Fagus dominated forests is enabled by the local mild mountain climate with average temperatures in July about 17 °C and average 
annual precipitation between 650–850 mm (cf. Tarábek 1980). 

3. METHODS 
Field research was carried out in June and July 2001. 
Thirty-two relevés were made following standard 
methods of the Zürich-Monpellier (Z-M) approach 
(Braun-Blanquet 1964) using the estimation abundance/dominancescaleaccordingtoBarkman&al. 
(1964). Relevés were stored and processed by the 
TURBOVEG program package (Hennekens 1996a) 
and MEGATAB program (Hennekens 1996b). 
For the numerical classification, TWINSPAN (Hill 
1979) program was used. For the indirect gradient 
analysis (Principal Components Analysis – PCA), 
CANOCO program (ter Braak & Šmilauer 1998) 
was used. Cover values of herb layer species were 
transformed into three levels: r, + = 1; 1, 2A, 2B = 
2; 3, 4, 5 = 3. The same downweighting of cover values was used by the TWINSPAN program, to make 
ordination and numeric classification closer to the 
floristic approach of the Z-M school. 

From each relevé plot, one soil sample of the 
top 10 cm of A horizon was taken. Soil reaction in 
H2O and KCl was measured; humus content was 
calculated according to the total organic carbon 
content (Tyurin’s method). Analyses were done in 
the laboratories of the Department of Forest Environment at the Technical University in Zvolen. 
Mean values for individual communities are presented in Fig. 2. 

Syntaxonomical units used are mostly sensu 
Mucina & al. (1993) or Moravec & al. (2000). Full 
names of syntaxa (including authors and the year 
of description or validation) are presented in the 
“Survey of vegetation units” or at the first mention 
in the text. The nomenclature of plant taxa follows 
Marhold & Hindák (1998). 

Determination of diagnostic species (cf. Whit-
taker 1962, Moravec 1994), dominants and prevailing species was based only on our relevé set 
presented in Table 1. We considered species with 
absolute cover of more than 25 % (3, 4, 5 degree of 
Braun-Blanquet scale) to be dominants and those 

62 


KAROL UJHÁZY, RICHARD HRIVNÁK, EVA BELANOVÁ, BLAŽENA BENČAŤOVÁ: THE BEECH FOREST VEGETATION OF THE CEROVÁ VRCHOVINA MTS. 

.......
.


..........
...........
Figure 2: Mean values of soil reaction and humus content of topsoil (A – Roso pendulinae-Tilietum cordatae, B – Athyrium filix-femina-Fagus sylvatica community, C – Mercurialis 
perennis-Fagus sylvatica community, D – Melico-Fagetum, 
E – Asperulo-Fagetum, F – Dentario bulbiferae-Fagetum, 
G – Carici pilosae-Fagetum, H – Luzulo-Fagetum) 

Slika 2: Srednje vrednosti pH in vsebnosti organske snovi v 
zgornji plasti tal (A – Roso pendulinae-Tilietum cordatae, 
B – združba Athyriumfilix-femina-Fagussylvatica, C – združba Mercurialis perennis-Fagus sylvatica, D – Melico-Fagetum, E – Asperulo-Fagetum, F – Dentario bulbiferae-
Fagetum, G – Carici pilosae-Fagetum, H – Luzulo-Fagetum) 

with more than 25 % of total herb layer cover to 
be prevailing species. Species that occur in more 
than 60 % of relevés of the current syntaxon were 
considered constant species. 

Within the characteristics of particular communities, the following abbreviations were used: ass. 

– association, art. – article of the Code of phytosociological nomenclature (Weber & al. 2000), c – 
constant species, corr. – corexit, dom. – dominant 
and prevailing species, dif. – differential species, E
– moss layer, E1 – herb layer, E2 – shrub layer, E
– tree layer, em. – emendavid, lok. – local, rel(s) 
– relevé(s), s. lat. – sensu lato. In Table 1 cover values 2a and 2b are presented as A and B. In the first 
column of Table 1, the following abbreviations are 
used: ca – Carpinion Issler 1931, cf – Cephalanthero-
Fagenion R.Tx. in Oberd. et R.Tx. 1958, fs – Fagion 
sylvaticae, Fs – Fagetalia sylvaticae, lf – Luzulo-Fagion, 
Pr – Prunetalia R.Tx. 1952, QF – Querco-Fagetea, Qp 
– Quercetalia pubescentis Klika 1933, QR – Quercetea 
robori-petraeae Br.-Bl. et R.Tx. ex Oberd. 1957, Qr 
– Quercetalia roboris R.Tx. 1931, ta – Tilio-Acerion. 
4. RESULTS 
4.1 Survey of vegetation units 
Querco-Fagetea Br.-Bl. et Vlieger in Vlieger 1937 
Fagetalia Pawłowski in Pawłowski, Sokołowski et 
Wallisch 1928 

Tilio-Acerion Klika 1955 em. Husová in Moravec et al. 1982 
Roso pendulinae-Tilietum cordatae Csiky et 
al. 2001 
Athyrium filix-femina-Fagus sylvatica community 
Mercurialis perennis-Fagus sylvatica community 
Dryopteris filix-mas-Fagus sylvatica community 
Fagion Luquet 1926 
Eu-Fagenion Oberd. 1957 em. R.Tx. in 
Oberd. et R.Tx. 1958 

Melico-Fagetum Seibert 1954 
Asperulo-Fagetum Souggnez et Thill 1959 
Dentario bulbiferae-Fagetum Zlatník 1935 
Carici pilosae-Fagetum Oberdorfer 1957 

Luzulo-Fagion Lohmeyer et R.Tx. in R.Tx. 
1954 
Luzulo-Fagetum Meusel 1937 

4.2 Characteristics of particular 
communities
According to our data set and its analyses (Table 1, 
Fig. 4) we could differentiate and characterise nine 
communities within three alliances of the Fagetalia 
order. 

Roso pendulinae-Tilietum cordatae 
(Table 1, rels 1–2; 
community A) 

Diagnostic species: Carpinus betulus (E3, c), Dryopteris filix-mas (c), Euonymus verrucosus (E, c, dif.), 

01

Fagus sylvatica (E, c, dom.), Hypnum cupressiforme 

33

(c, dom.), Isothecium myurum (dom.), Tilia platyphyllos (E3, c, dom.; E1, c, dif.). 

This community was described recently just 
from the Cerová vrchovina Mts. (Csiky & al. 2001). 
Our relevés document this community from two 
other localities. 

The community is well differentiated by species 
composition and habitat character. In the tree layer, coppices of Tillia platyphyllos dominate, in some 
places Fagus sylvatica prevails. Tilia platyphyllos frequently replaces T. cordata in stands of this com

63 


HACQUETIA 3/1 • 2004 

Table 1: Tabular differentiation of communities with Fagus sylvatica in the Cerová Vrchovina Mts. (A – Roso pendulinae-
Tilietum cordatae, B – Athyrium filix-femina-Fagus sylvatica community, C – Mercurialis perennis-Fagus sylvatica community, 
D – Melico-Fagetum, E – Asperulo-Fagetum, F – Dentario bulbiferae-Fagetum, G – Carici pilosae-Fagetum, H – Luzulo-Fagetum) 

Tabela 1: Tabelarični prikaz razlik združb z vrsto Fagus sylvatica v hribovju Cerová Vrchovina (A – Roso pendulinae-Tilietum 
cordatae, B – združba Athyrium filix-femina-Fagus sylvatica, C – združba Mercurialis perennis-Fagus sylvatica, D – Melico-
Fagetum, E – Asperulo-Fagetum, F – Dentario bulbiferae-Fagetum, G – Carici pilosae-Fagetum, H – Luzulo-Fagetum) 

Relevé number: 1 2 
Number of species in 

93

herb layer: 
Community: 

A 
Tree layer 

B C D E F G H 
fs Fagus sylvatica +5 
ta Tilia platyphyllos 

41 
ta Acer platanoides 

B. 
ta Ulmus glabra 
.. 
ta Acer pseudoplatanus 

.. 
ca Carpinus betulus 

+1 
Quercus petraea agg. .. 
ca Cerasus avium .. 
QF Acer campestre .. 
ta Tilia cordata .. 

Shrub layer 

QF Corylus avellana A. 
ta Tilia platyphyllos 

A. 
fs Fagus sylvatica .+ 
ta Acer platanoides .. 
Diagnostic species 

Qp, Pr Euonymus verrucosus (E1) 

r+ 
r+ 
Stachys sylvatica .. 
Urtica dioica .. 
Athyrium filix-femina .. 
Circaea lutetiana .. 
Sambucus nigra .. 
Clematis vitalba .. 
ta Chelidonium majus .. 
fs, Fs Mercurialis perennis .. 
fs, Fs Galeobdolon luteum .. 
fs, Fs Polygonatum multiflorum .. 
fs, Fs Isopyrum thalictroides .. 
fs, Fs Glechoma hirsuta .. 
Fs Melica uniflora .. 
fs, Fs Dentaria bulbifera .. 
ca Carex pilosa .. 
lf, QR Veronica officinalis .. 
lf, QR Hieracium lachenalii .. 
QF, Qp, cf Campanula persicifolia .. 
Qp, QF Hylotelephium maximum .. 
ca, Qr Hieracium sabaudum .. 
Veronica chamaedrys .. 
ca Galium schultesii .. 
QF Poa nemoralis .. 
lf, Qr Luzula luzuloides .. 
Pohlia nutans .. 
Hieracium murorum .. 
fs, Fs Campanula trachelium .. 

ta Tilia platyphyllos (E1) 


34567 

2414 9 1614 

54454 
.A. . . 
.1. . . 
.1. . . 
. .1.3 
..... 
..... 
..... 
..... 
..... 

. . . .+ 
.1. . . 

+1. . . 
.+. . . 
..... 

..... 

r ++ 

.. 

. 1+ r+ 

AA1 1 + 

1AA11 

++ . 1+ 

+++. r 

...rr 

..... 

..... 

..... 

..... 

..... 

....r 

..... 

..... 

..... 

..... 

..... 

..... 

..... 

..... 

..... 

r.... 
r.... 
..... 
..... 
..... 
8 9 1011 1213141516171819202122232425262728293031 
7 201818162528282215171311 7 10 9 12 5 9 1633272618 

35A41455B555555545554545 
. .B.A 

................... 
. +313 

. . . . . . . . . . .A. . . . . . . 
. .B. . 

..................
.


................... 
+.1B11. .3A. . .++++. . .+. .+ 

3.1.+ 
1. .+11. .3+. . . .11. . . . r . . . 
. . . . . . . . . . . . .1. .1. . . . . . . 
. . .1. . . . . . . . . . . .+. . . . . . . 
. . . . .+. . . . . . . . . . . . . . . . .+ 
........................ 
. .+.+ 

................... 
. .+.+.++.+1A. .1.+++++.++ 
. .+. . . . . . . . . . . . . . . . . . . . . 

. r . .+. . .+. . . . . . . . . . . . . . . 
. . . . . . . . . . . . . . r .+. . . . . . . 
. . . . . . . . . . . . . . . . . . .+. . . . 
. . .+.+. . . . . . . . . . . . . . . . . . 
. . . . . . . . . . . . . . . r . . .1. . . . 
. . .+.+. .+. .++. .+. .+1. . . . 
.rr...r.r...rr..r....... 
. . . . . . r+. .++. . . . . . .+r . . . 
.A++. r+ 

. . .+. . . . . . . . . . . . . 
3B333B4 

1. .+. .1. . . . . . . . .+ 
. .A+1.A 
................
.


... 

.+. . . . . . . . . . . .+. . . 
... 

r ++ 

. ++ 

.................. 
... 

11 . 
.................. 
. . . r+.+ 

343 

.............. 
. . . .11+. . . . . r . 

+11 
....... 
. . . . . .+. . . . . . . . .1 

1. . . 
.................... 
B43 

. 

...................
.


+r . 
A+ . 

. 
.................... 

+.+ 

. 
.................... 

r.r 
. 
.................... 

+. .+ 
.r.................. 

++ r+ 
. . . . . . . . . .+. . . . . . . . . 

+r A1 
. . . . . . . . . .1. . . . . . . . . 
3BA+ 
.................... 

+1 rA 
.................... 
1 . 1+ 
..................... 
++B 
..................... 

. ++ 


64 


KAROL UJHÁZY, RICHARD HRIVNÁK, EVA BELANOVÁ, BLAŽENA BENČAŤOVÁ: THE BEECH FOREST VEGETATION OF THE CEROVÁ VRCHOVINA MTS. 

Relevé number: 1 2 3 4 5 6 7 8 9 1011 1213141516171819202122232425262728293031 

Constant species 

fs, Fs Dryopteris filix-mas 11AA1++.1++Br+. .+1.+r . r r+.+. r++ 
fs Fagus sylvatica . .+. .+r r . . . . .+++++++r+.++.+++r . 
fs, Fs Mycelis muralis . .1+.++.+r . . r++r r+.+. r . . . .++1++ 

Herb layer 

fs, Fs Geranium robertianum 

. . .+. . . . . . . . .+. . . 
ta Acer pseudoplatanus 

r .++1.+. .+. r1+ 
. . r .+.++. .1r+. 

..............r.. 
fs Actaea spicata 

. . . .++. . . .+. . . 

. . .1. . . . . . . . . . . . . 
Fs Scrophularia nodosa 

..r...r.r..... 

................
.


r . . . . . . . .+. . . .
Asplenium trichomanes 

................. 
ta Acer platanoides +. .+. . . .++1. r1+. . .+. . r+1. . . .+. r 
QF Acer campestre . . r . .+.+. . r . . .+1. .1. . r . r .+++. . . 
fs, Fs Viola reichenbachiana . .1+.+.1. .+.+++B++A1+. . . .+11. . . 
fs, Fs Galium odoratum . . r . . . .+BABAA+BA.1+++. .1. .++r r . 
Moehringia trinervia . . r . . . . .++. . r+. . . . . . . . . . . . .+++. 
ta Alliaria petiolata . . r . . . r+. .111+++. . r . . . . . . . . r r . . 
Galium aparine . . . . . r . . r .++.1+++. . r . . . . . . .++r+ 
Fallopia convolvulus . . . . . . . . r . . .++1+. . .+. . . . . . .+++. 
Lactuca serriola ........+r. +rrr...........r..+ 
fs, Fs Lathyrus vernus . . . . . . . . . . .+r r+.++r . . r+. .+++rA+ 
ca, cf (Qp) Melittis melissophyllum . . . . . . . . . . . . .++. . . . . . . . . . . . r . r . 
ca, Qp Lathyrus niger . . . . . . . . . . . . .++. . . . . . . . . . . . . . . . 
ca Cerasus avium . . . . . r . . r . . . r+r+r . . . r . .+. r . . r . . 
Rosa canina agg. .............+rr............... 
Epilobium montanum ........r....rrr...........r... 
ca Carpinus betulus . . . . . . . . . . r . . . .++. . . . . . . . . . . . . . 
fs, Fs Pulmonaria obscura . . .+. . . . . . . . r+++. r+. . . . . . r . . . . . 
fs, Fs Tithymalus amygdaloides . . . . . . . . . . . . .+.++++. .+r r . . .+. . . 
Quercus petraea agg. .....+..........+....rr..+..rr. 
fs, Fs Carex digitata . . r . . . . . . . . . . . . . . . . . .+. .+++.+.+ 
Campanula rapunculoides +. . . . . . .+r . . . .A. . r . . . . . . . . .A.11 
Cruciata glabra . . . . . . . . . . . . . .+. . . . . . . . . . . .1+. . 
Ajuga reptans . .+. . . . . . . . . . . . . . r . . . . . . . . .++. . 
Polypodium vulgare +. . . . . . . .+. . . . . . . . . . . . . . . . . . r+.

 -species with occasional occurence 
fs, Fs Neottia nidus-avis r...............r.............
.
Rubus hirtus s.lat. ..++...................r..+...
.
Qp Quercus cerris . . r . . . . .+. . . . . . . . . . . . . . . . . . . . r 
.
QF Hedera helix . . . . .+. . . . . . . . . . . . . . . . . .+. . . . . 
.
Robinia pseudoacacia . . . . . r . . . . . .+. r . . . .+. . . . . . . . . r 
.
Cardaminopsis arenosa . . . . . . . . r+. . . . . . . . . . . . . . . . . . . .1
Lapsana communis ........r...r................r.
ca Dactylis glomerata . . . . . . . . . . . . . .+. . . . . . . . . . . . .+. 
.
Fs Carex sylvatica . . . . . . . . . . . . . . .+. . . . . . . . . .+. . . 
.
Symphytum tuberosum . . . . . . . . . . . . . . . .+. r . . . . . . . . . . r 
.


Moss layer 

Isothecium myurum B. . . . . . . . . .+. . . . . . . . . . . . . . . . . . . 
Hypnum cupressiforme 3A.A. . . . . . .A.+.+.+. . .++. .+.+. . . 
Paraleucobryum longifolium .+.+. . . . . . . . . . . . . . . . .+. . . . . . . . . 
Schistidium apocarpum .+.+. . . . . . .1. . . . . . . . .+. . . . . . . . . 
Brachythecium velutinum .+. . .A. . . . . . . . . .++. . . . . . .+.1A+. 
Atrichum undulatum . . .+. . . . . . . . . . . . . . . . . . . . .+.+. . . 
Dicranella heteromalla . . . . .+. . . . . . . . . . . . . . . . . . .+. .+. . 
Plagiothecium cavifolium . . . . . . . . .1. . . . . . . . . . . . . . . . . . . .B 
Pylaisia polyantha . . . . . . . . .+. . . . . . . . . . . . .+. . . . . . . 
Dicranum scoparium . . . . . . . . . . .+. . . . . . . . .+. . . . . . . .+ 

65 


HACQUETIA 3/1 • 2004 

munity (cf. Csiky & al. 2001). In the shrub layer, 
especially Coryllus avellana, Euonymus verrucosus and 
Tilia platyphyllos occur. From the constant species 
presented by Csiky & al. (2001), we found only 
Dryopteris filix-mas, Euonymus verrucosus, Geranium 
robertianum and Polypodium vulgare. The herb layer 
features very low cover, which fully complies with 
the assertions of the above mentioned authors. On 
the other hand, the high cover features moss layer, 
with Hypnum cupressiforme as the most frequent and 
dominant species. 

Roso pendulinae-Tilietum cordatae is a typical block 
forest community. Basalt boulders covered more 
than 60 % of the plots. Slopes were steep, exposed 
to the north. On the first relevé plot, soil reaction 
was acid, on the second it was neutral. The share 
of humus and total nitrogen was very high. These 
values are several times higher in comparison with 
other Tilio-Acerion communities in this region (see 
Fig. 2). 

The following three communities are of transitional character between Tilio-Acerion and Fagion 
alliances. At the moment, they have been classified 
into the Tilio-Acerion alliance according to floristic 
composition and some ecological conditions. 

Athyrium filix-femina-Fagus sylvatica community 
(Table 1, rels 3–7; B) 

Diagnostic species: Acer pseudoplatanus (E3, 
dom.), Athyrium filix-femina (c, dif.), Brachythecium 
velutinum (dom.), Circaea lutetiana (c), Clematis vitalba (c), Dryopteris filix-mas (c), Fagus sylvatica (E3, 
c, dom.), Geranium robertianum (c), Hypnum cupressiforme (dom.), Mycelis muralis (c), Sambucus nigra 
(c), Stachys sylvatica (dif.), Urtica dioica (c, dif.). 

Fagus sylvatica is a dominant species of the tree 
layer in all relevés, Acer pseudoplatanus, A. platanoides, Tilia platyphyllos and Ulmus glabra make the 
admixture. The shrub layer is weakly developed 
in commercial forests. Apart from young trees, we 
recorded only Corylus avellana and Sambucus nigra. 
The herb layer is represented by 15 species per 
relevé on average, with the range of cover between 

(2) 8–25 %. It is characterised by a high cover of 
ferns. The species Athyrium filix-femina and Circaea 
lutetiana have their distinct ecological optimum in 
this community within beech-dominated forests in 
the Cerová vrchovina Mts. These two species along 
with Dryopteris filix-mas are the most dominant ones 
in the herb layer. The other species have only low 
cover values. 
The community occurs on rocky, steep and 
shady endings (foots) of slopes above moist val

leys, or directly in the side valley branches. Along 
valleys, it descends to low altitudes (the lowest occurrence at 335 m a. s. l.). The soils developed on 
various bedrock (andesite, sandstone, basalt) are 
slightly acid, rarely acid. They show relatively high 
values of humus content (Fig. 2) in comparison 
with the next communities. However, the most important ecological factor of these habitats seems to 
be a high aerial moisture, which is reflected in the 
increased dominance of ferns. 

Mercurialis perennis-Fagus sylvatica community 
(Table 1, rels 8–14; C) 

Diagnostic species: Acer platanoides (E3, dom.; 
E1, c), Acer pseudoplatanus (E3, dom.), Alliaria petiolata (c), Carpinus betulus (E3, c, dom.), Dryopteris filix-mas (c), Fagus sylvatica (E3, c, dom.), Galeobdolon 
luteum (c, dif.), Galium odoratum (c, dom.), Glechoma hirsuta (dif.), Hypnum cupressiforme (dom.), Chelidonium majus (c, dif.), Isopyrum thalictroides (dif.), 
Isothecium myosuroides (dom.), Mercurialis perennis 
(c, dif., dom.), Polygonatum multiflorum (dif.), Tilia 
platyphyllos (E3, dom.), Ulmus glabra (E3, dom.). 

The tree layer of this heminitrophilous community is relatively species-rich. It is formed mainly by 
Fagus sylvatica with a variable admixture of nitrophilous tree species (Acer platanoides, A. pseudoplatanus, 
Tilia platyphyllos, Ulmus glabra) and Carpinus betulus 
(locally also with Quercus petraea agg.). The shrub 
layer was weakly developed under the closed canopy of trees. We found about 19 species of herb layer 
on the relevé plots. A strong dominant of the layer 
is Mercurialis perennis, frequently accompanied by 
Galium odoratum and locally also by Galeobdolon luteum. In some cases, more abundant were Dryopteris 
filix-mas, Alliaria petiolata, Glechoma hirsuta, Dentaria bulbifera and Chelidonium majus. The most frequently naturally regenerating tree species are Acer 
platanoides and A. pseudoplatanus. We recorded the 
cover of herb layer of (25–35) 50–80 %. 

We found this community at relatively higher 
altitudes (470–700 m a. s. l.), mostly about 550 m. It 
grows from moderate to very steep slopes, exposed 
to the northeast, north and northwest. The soil reaction varies from acid to slightly acid. The average pH value is higher than in mesotrophic beech 
forests (Fig. 2). 

The community is well distinguished from the 
syntaxa of mesotrophic beech forests by the occurrence of several nitrophilous species (Chelidonium majus, Geranium robertianum). Relevés No. 10 
and 12 (Table 1) on the more bouldery soils with 
a higher occurrence of nitrophilous tree species 

66 


KAROL UJHÁZY, RICHARD HRIVNÁK, EVA BELANOVÁ, BLAŽENA BENČAŤOVÁ: THE BEECH FOREST VEGETATION OF THE CEROVÁ VRCHOVINA MTS. 

are close to the Mercuriali-Fraxinetum (Klika 1942) 
Husová in Moravec et al. 1982 association. Klika 
(1942) described this association as “Acereto-Fagetum carpaticum”. The original name had to be rejected according to the Code of Phytosociological 
nomenclature (art. 34). Husová (2000) as well as 
Fajmonová (1984) consider this Mercuriali-Fraxinetum as a mountain forest community on rocky or 
bouldery screes. However, the communities with 
Mercurialis perennis in Cerová vrchovina in most cases (relevés Nr. 8, 9, 11, 13 in Table 1) were found 
on cambisols with a normally developed A horizon, 
with the surface stoniness below 25 %. Their typical feature is a high content of small soil skeleton 
in the topsoil. 

Species composition is different as well. Unlike 
the Mercuriali-Fraxinetum, several diagnostic species 
(such as Dentaria eneaphyllos, Hordelymus europaeus, 
Fraxinus excelsior) are missing in our communities. 
Therefore, we named it the Mercurialis perennis-Fagus sylvatica community. According to the species 
composition, we ranked it into Tilio-Acerion alliance, however it has transitional character towards 
the Fagion alliance. 

Dryopteris filix-mas-Fagus sylvatica community 

One specific community we recorded on andesites near the top of Karanč (725 m a. s. l.) on a 
steep rocky slope (about 50 % of soil skeleton in 
the topsoil). 

Šiatorská Bukovinka, SE from the village, Karanč 
(725 m a. s. l.), about 250 m to the NW from the 
top of the hill; beech forest on a steep slightly concave slope close below the ridge; rocky (andesite) 
soil covered with thick litter layer; longitude: 19° 
47,4639’, latitude: 48° 9,61284’; age: 60–80 years; 
slope: 31°; aspect: 350°; relevé area: 400 m2; total 
cover: 90 %; E3: 90 %; E1: 20 %; height of the tree 
layer: 21 m; 12.6.2001; B. Benčaťová, K. Ujházy; 
field number: 21/01, (relevé 32 in Figure 4). 

E3: Fagus sylvatica 5, Tilia cordata +. 

E1: Dryopteris filix-mas 2b, Acer pseudoplatanus +, 
Chelidonium majus +, Robinia pseudoacacia +, Cerasus avium r, Fallopia convolvulus r, Geranium robertianum r, Moehringia trinervia r, Prenathes purpurea r, 
Urtica dioica r. 

It is a beech forest with the Tilia cordata admixture. The poorly developed herb layer has a heminitrophilous character with the prevalence of Dryopteris filix-mas. Mountain character is indicated by 
the occurrence of Prenanthes purpurea. 

According to the occurrence of species such as 
Chelidonium majus, Urtica dioica, Geranium robertianum, we decided to rank this community to the 
Tilio-Acerion alliance. Similar communities were recorded in the Ostrôžky Mts. in the southeast part 
of the volcanic Slovenské stredohorie mountain region (Benčaťová, Hrivnák & Ujházy in litt.). A more 
exact classification will be possible after the comparison of a larger data set (from volcanic regions 
of southern Slovakia and northern Hungary). 

Melico-Fagetum (Table 1, rels 15–17; D) 

Diagnostic species: Alliaria petiolata (c), Carpinus 
betulus (E3, dom.; E1, c), Cerasus avium (E1, c), Epilobium montanum (c), Fagus sylvatica (E3, c, dom.; E2, 
E1, c), Fallopia convolvulus (c), Galium aparine (c), 
Galiumodoratum (c,dom.),Lactucaserriola (c), Lathyrus vernus (c), Melica uniflora (c, dif., dom.), Mycelis 
muralis (c), Pulmonaria obscura (c), Quercus petraea 
agg. (E3, dom.), Rosa canina agg. (E1, c), Tithymalus 
amygdaloides (c), Viola reichenbachiana (c). 

Melico-Fagetum is a medium species-rich community (23 taxa per relevé on average). Fagus sylvatica 
dominates in the tree layer. At lower altitudes, it 
is accompanied by Carpinus betulus and Quercus 
petraea agg. Shrubs occur rarely or are missing. The 
herb layer has a high cover value (more than 50 
%). Along with the general dominant species Melica uniflora, only two species (Galium odoratum and 
Viola reichenbachiana) have a high cover in some 
patches as well. Other species occur only with low 
cover values. Apart from the typical beech forest 
species, some species of Carpinion alliance (Acer 
campestre, Crataegus spec. div., Melittis melisophyllum, 
Rosa canina agg.) also grow here. 

We detected the stands of Melico-Fagetum at altitudes between 400–500 m a. s. l. on even, slightly 
convex or concave slopes with an inclination of less 
than 15° without soil surface stoniness. Gravelly 
topsoil is strongly rooted. Soil reaction is slightly 
acid and acid, values of humus content are similar to those of the other Eu-Fagenion communities 
(Fig. 2). 

The so-called “Fagetum nudum” communities are 
the most frequent types of beech forests in this region. Many of them are almost without herb layer. 
Types with a developed herb layer can be divided 
into the next two associations: 

Asperulo-Fagetum (Table 1, rels 18–21; E) 

Diagnostic species: Dryopteris filix-mas (c), Fagus 
sylvatica (E3, c, dom.; E1, c), Galium odoratum (c), 
Viola reichenbachiana (c). 

67 


HACQUETIA 3/1 • 2004 

Fagus sylvatica is the absolute dominant of the 
tree layer. Only in the heminitrophilous variant 
with Mercurialis perennis (transition to the Mercurialis 
perennis-Fagus sylvatica community), did we record 
the admixture of Carpinus betulus and Cerasus avium. The shrub layer is formed by locally frequent 
beech. Apart from several mesotrophic species of 
the Fagion alliance or Fagetalia order, respectively 
(mostly Viola reichenbachiana and Galium odoratum), 
also some heminitrophilous species (such as Circaea lutetiana, Mercurialis perennis) with low cover 
values occur in the herb layer. The community has 
no differential species. 

In contrast to the Dentario bulbiferae-Fagetum, we 
found Asperulo-Fagetum stands at higher altitudes 
(460–600 m a. s. l.). The Carpinion alliance species 
are missing on these habitats. The soils are cambisols with relatively higher humus content and 
higher pH values in the topsoil (Fig. 2). 

We suppose that this community represents 
both qualitatively and quantitatively, the substantially impoverished variant of the low-herb flowery 
Carpathian beech forests on (possibly beyond) the 
southern border of their occurrence area. From the 
syntaxonomical point of view they are most close to 
the Central-European association Asperulo-Fagetum 
according to Wallnöfer & al. (1993). Quite similar 
communities were described by Moravec (1979) 
from the basalt Doupovské hory Mts. in northwest Bohemia as the Violo reichenbachianae-Fagetum 
Moravec 1979. The most important characteristic 
species of these communities – Hordelymus europaeus – is missing in the species-poor communities of 
the Cerová vrchovina Mts. 

Dentario bulbiferae-Fagetum 
(Table 1, rels 22–24; F) 

Diagnostic species: Acer campestre (E1, c), A. platanoides (E1, c), Carpinus betulus (E3, c), Dentaria bulbifera (c, dif.), Dryopteris filix-mas (c), Fagus sylvatica 
(E3, c, dom.; E2, E1, c), Lathyrus vernus (c), Quercus 
petraea (E3, E1, c), Tithymalus amygdaloides (c). 

We found this community at lower altitudes 
(370–430 m a. s. l.). From other studied forest communities of this territory, it is differentiated negatively (there are missing several diagnostic species 
of the Fagion alliance, or Fagetalia order, e. g. Galium odoratum, Viola reichenbachiana, Mycelis muralis). 
The only diagnostic (and also differential) species 
is Dentaria bulbifera. 

In the beech-dominated tree layer, also Quercus 
petraea agg. with Carpinus betulus occur, rarely Acer 
platanoides, Cerasus avium and Acer campestre. In the 
weakly developed shrub layer of the commercial 

stands, only beech occurs. The herb layer is very 
poor. Only Dentaria bulbifera is more frequent in the 
vernal aspect. Other (mostly mesotrophic) species 
have cover below 1 % on the relevé area. On some 
places natural regeneration of tree species occurs. 

The stands grow on even or slightly convex slopes without or with only a small stoniness on the 
soil surface. Normally developed soil exhibits acid 
reaction. The average values of humus content are 
similar to those in the Mercurialis perennis-Fagus sylvatica community and in Luzulo-Fagetum (Fig. 2). 

Carici pilosae-Fagetum (Table 1, rels 25–27; G) 

Diagnostic species: Acer campestre (E1, c), Carex 
digitata (c), Carex pilosa (c, dif., dom.), Dryopteris filix-mas (c), Fagus sylvatica (E3, c, dom.; E2, E1, c), 
Lathyrus vernus (c), Viola reichenbachiana (c) 

The tree layer of this community in the Cerová 
vrchovina Mts. was completely dominated by Fagus 
sylvatica. The shrub layer was weakly developed, 
formed of young individuals of beech, which is 
also most frequently regenerating. It is a species-
poor community (with only 10 species per relevé 
on average) with the Carex pilosa dominance in the 
herb layer. Probably under the influence of the drier climate of the region, the herb layer has lower 
cover than communities in the Carpaticum floristic 
region. 

The stands occur on moderate slopes. The soil 
is light grey brown, acid with higher sand content. 
Among other studied communities, this community has the lowest average values of soil reaction 
and humus content (Fig. 2). 

Luzulo-Fagetum (Table 1, rels 28–31; H) 

Diagnostic species: Brachythecium velutinum (c), 
Campanula persicifolia (dif.), C. rapunculoides (c), 

C. trachelium (dif.), Dryopteris filix-mas (c), Fagus sylvatica (E3, dom., c; E2, E1, c), Fallopia convolvulus 
(c), Galium aparine (c), G. odoratum (c), G. schultesii (c, dif., dom.), Hieracium lachenalii (dom., dif.), 
H. murorum (c, dif., dom.), H. sabaudum (dif.), 
Hylotelephium maximum (dif.), Lathyrus vernus (c, 
dom.), Luzula luzuloides (c, dif., dom.), Moehringia 
trinervia (c), Mycelis muralis (c), Plagiothecium cavifolium (dom.), Poa nemoralis (c, dif., dom.), Pohlia nu-
tans (c, dif.), Veronica chamaedrys (c, dif.), Veronica 
officinalis (dif.). 
Fagus sylvatica is the only dominant species of 
the tree layer. Carpinus betulus forms the admixture 
in some places. Only young individuals of Fagus 
sylvatica, or rarely other tree species, occur in the 
shrub layer. On the other hand, on average, 26 spe

68 


KAROL UJHÁZY, RICHARD HRIVNÁK, EVA BELANOVÁ, BLAŽENA BENČAŤOVÁ: THE BEECH FOREST VEGETATION OF THE CEROVÁ VRCHOVINA MTS. 

cies occur in the herb layer, and also the moss layer 
is relatively rich. Consequently, it is the most species rich community among other Fagus dominated 
communities of the region. The community is well 
differentiated through several oligotrophic species 
(Luzula luzuloides, Veronica officinalis or Hieracium 
spec. div.) and some mesotrophic Carpinion alliance diagnostic species (Galium schultesii, Veronica 
chamaedrys). However, also a lot of Fagetalia order 
or Fagion alliance diagnostic species exhibit a high 
constancy. 

We recorded this community at altitudes about 
600 m a. s. l. (570–630) on the northwest aspect. 
The communities are developed on steep convex slopes or side ridges. Soils are shallow, rocky 
and easily drying, with acid or less frequently with 
slightly acid reaction (Fig. 2). 

Only the last relevé (Table 1, rel. 31) on rhyodacite (probably) tuffs has a distinctively oligotrophic 
character. This community can be classified as Luzulo-Fagetum luzuletosum albidae (Tüxen 1937) Hartmann 1953 em. Moravcová-Husová 1964. 

The other three relevés (Table 1, rels 28–30) 
with Poa nemoralis dominance or subdominance 
contain many mesotrophic diagnostic species of 
Fagion alliance. Phytocoenoses of beech forest with 
Poa nemoralis dominance at lower altitudes were 
described by Moravcová-Husová (1964) as Luzulo-
Fagetum poetosum nemoralis Moravcová-Husová 1964. 
Moravec (2000) later classified these communities 
within the Luzulo-Fagetum luzuletosum albidae subassociation. 

5. DISCUSSION AND CONCLUSIONS 
Tabular synthesis according to TWINSPAN analysis (Table 1) and indirect gradient analysis (PCA) 
yielded quite similar results, if the herb layer species cover values were transformed to the three 
levels (Fig. 4). Presence/absence data themselves 
were not sufficient for precise classification of 
these communities, however, floristic composition 
is the most important classification criterion of the 
Zürich-Montpellier approach. 

By means of the above mentioned methods, we 
could specify nine plant communities with Fagus 
sylvatica ranked into three alliances in the Cerová 
vrchovina Mts. 

Heminitrophilous and nitrophilous communities of the Tilio-Acerion alliance represent four units. 
They occur in side valley branches or on slopes 
with increased surface stoniness or high content of 

soil skeleton in the topsoil. These rocky soils are 
mostly slightly acid, less frequently acid and rarely neutral. Average pH values and percentages of 
humus content are higher than in beech forests 
of the Fagion or Luzulo-Fagion alliances (Fig. 2). In 
adition to beech, which is a common dominant 
(even in scree forests), also nitrophilous species of 
the genera Tilia, Acer, Ulmus and Carpinus betulus 
are frequent in the tree layer. However, some pure 
beech stands were classified within the Tilio-Acerion 
alliance according to herb layer diagnostic species. 
Cover of the herb layer varies according to stoniness. The moss layer is best developed on boulder 
fields. Here it reaches a higher cover than the herb 
layer and also the highest cover values of moss layer 
within all studied communities (Fig. 3). 

..
. 

..
. 

..
. 

..
. 

..
. 

..
. 

..
. 

.
.
.......
.


...
...
Figure 3: Mean cover values of herb and moss layer in individual communities (A – Roso pendulinae-Tilietum cordatae, 
B – Athyrium filix-femina-Fagus sylvatica community, C – Mer-
curialis perennis-Fagus sylvatica community, D – Melico-Fagetum, E – Asperulo-Fagetum, F – Dentario bulbiferae-Fagetum, 
G – Carici pilosae-Fagetum, H – Luzulo-Fagetum) 

Slika 3: Srednja pokrovna vrednost zeliščne in mahovne plasti 
v posamezni združbi (A – Roso pendulinae-Tilietum cordatae, 
B – združba Athyrium filix-femina-Fagussylvatica,C –združba 
Mercurialis perennis-Fagus sylvatica, D – Melico-Fagetum, 
E – Asperulo-Fagetum, F – Dentario bulbiferae-Fagetum, 
G – Carici pilosae-Fagetum, H – Luzulo-Fagetum) 

Mesotrophic beech forests of the Fagion alliance 
represent five communities. They occupy most frequently even slopes and, unlike the Tilio-Acerion 
communities, they grow on habitats without (or 
with a small percentage of) surface stoniness. The 
single dominant of the tree layer is beech. At lower 
altitudes, it is accompanied by oaks and hornbeam. 
“Scree” species of the genera Tilia, Acer and Ulmus 

69 


HACQUETIA 3/1 • 2004 

are rare or completely missing. The cover of the 
herb layer is distinctively higher than the cover of 
the moss layer, which is frequently not developed. 

A special position is held by hemioligotrophic 
beech forests of convex sites with shallow soils with 
lower humus content (Fig. 2). These relatively species-rich (both herbs and mosses) communities, 
classified as Luzulo-Fagetum, are well floristically 
differentiated from the others. It has a transitional character between the Fagion and Luzulo-Fagion 
alliance, because strictly acidophilous herb species 
are missing there. 

Differentiating among of several communities was complicated because of the insufficient 
number of differential species, namely in the case 
of mesotrophic beech forests, which show a low 
species diversity and low cover values of herbs and 
mosses. Some of them could be differentiated only 
negatively. We also met some problems with ranking the particular communities into syntaxonomi-


.
...............................
....
.
..
..
..
..
..
.
..
..
.
..
..
..
..
..
.....
..
..
..
..
....
..
..
Figure 4: Ordination of relevés according to transformed herb 
layer species cover values using Principal Component Analysis. 
First ordination axis can be explained as nutrient/pH gradient 
(oligotrophic communities on the left side and heminitrophytic 
on the right). Relevés of individual communities are marked 
as: 
= 
– Roso pendulinae-Tilietum cordatae, 
. 
– Athyrium filix-femina-Fagus sylvatica community, 
. 
– Mercurialis perennis-Fagus sylvatica community, 
x 
– Dryopteris 
filix-mas-Fagus sylvatica community, 
. 
– Melico-Fagetum, 
# – Asperulo-Fagetum, 
. 
– Dentario bulbiferae-Fagetum, 
. – Carici pilosae-Fagetum, . – Luzulo-Fagetum 

Slika 4: Ordinacija popisov z metodo glavnih komponent 
(PCoA). Pokrovne vrednosti zeliščne plasti so transformirane. 
Prva ordinacijska os predstavlja gradient hranil in pH, (oligotrofne združbe na levi in zmerno nitrofilne na desni strani). 
Popisi posameznih združb so označeni: 
= 
– Roso pendulinae-
Tilietum cordatae, 
. 
– združba Athyrium filix-femina-Fagus sylvatica, 
. 
– združba Mercurialis perennis-Fagus sylvatica, x – združba Dryopteris filix-mas-Fagus sylvatica, 
. 
– Melico-Fagetum, # – Asperulo-Fagetum, 
. 
– Dentario bulbiferae-Fagetum, 
. 
– Carici pilosae-Fagetum, 
. – Luzulo-Fagetum 

cal units. This is discussed directly in the Results 
chapter. Generally, the communities shows floristical similarity with forest communities of the volcanic regions of the south of Central Slovakia, but 
they are significantly different (both in structure 
and species composition) from typical Carpathian 
communities, which were described from Slovakia. 
These syntaxonomical problems should be solved 
only by means of comparison of the larger data 
set from the volcanic regions of Matricum and Carpathicum regions and by cooperation between Slovak and Hungarian researchers. 

6. ACKNOWLEDGEMENTS 
We are grateful to Dr. R. Šoltés for mosses determination and Dr. A. Guttová for lichens determination. This paper was supported by the Slovak Grant 
Agency for Science (grants No. 1/7057/20 and 
1/0629/03). 

7. REFERENCES 
Balázs, P. (1996): Jelšiny Cerovej vrchoviny (fytocenologická charakteristika). Ochr. Prír. 14: 
29–39. 

Barkman, J. J., Doing, H. & Segal, S. (1964): Kritische Bemerkungen und Vorschläge zur quantitativen Vegetationsanalyse. Acta Bot. Neerl. 

13: 394–419. 
Braun-Blanquet, J. (1964): Pflanzensoziologie. 
Springer Verlag, Wien, 865 pp. 
Csiky J., Kevey B. & Borhidi A. (2001): Block forest 
(Roso pendulinae-Tilietum cordatae), a new forest 
community of the Carpathian basin (Cerová vrchovina, Slovakia). Acta Bot. Hung. 43 (1–2): 
95–125. 

Fajmonová, E. (1984): Mercuriali-Fraxinetum v severnej časti Bielych Karpát. Biológia 39: 101–104. 

Futák, J. (1966): Fytogeografické členenie Slovenska. – In: Futák, J. (ed.): Flóra Slovenska I. Veda, 
Bratislava, pp. 539–544. 

Hendrych, R. (1959): Florografický materiál z okolí 
Rimavské Soboty. Preslia 31: 187–207. 

Hendrych, R. (1963): Ad f loram dicionis oppidi 
Šafárikovo in Slovacia materies critica. Biol. 
Práce Slov. Akad. Vied 9(6): 1–63. 

Hendrych, R. (1968): Ad floram regionis Fiľakoviensis in Slovacia addenda critica. Acta Univ. 
Carol., Biol. 1967: 109–183. 

70 


KAROL UJHÁZY, RICHARD HRIVNÁK, EVA BELANOVÁ, BLAŽENA BENČAŤOVÁ: THE BEECH FOREST VEGETATION OF THE CEROVÁ VRCHOVINA MTS. 

Hennekens, S. M. (1996a): TURBO(VEG). Software package for input, processing, and presentation of phytosociological data (User’s guide). 
IBN-DLO, Wageningen et University of Lancaster, Lancaster, 52 pp. 

Hennekens, S. M. (1996b): MEGATAB a visual editor for phytosociological tables (version 1.0). 
Gielsen & Geurts, Ulft, 10 pp. 

Hill, M. O. (1979): TWINSPAN. A Fortran program 
for arranging multivariate data in an ordered 
two-way table by classification of the individuals 
and attributes. Cornell Univ., Ithaca, 52 pp. 

Holub, J. & Moravec, J. (1965): Floristische Materiale aus Hügellande Fiľakovská hornatina (Südslowakei). Biol. Práce Slov. Akad. Vied 11(6): 
1–91. 

Husová, M. (2000): Svaz: Tilio-Acerion. In: Moravec, 
J., Husová, M., Chytrý, M. & Neuhäuslová, M. 
(eds.): Hygrofilní, mezofilní a xerofilní opadavé lesy. Academia, Praha, pp. 115–140. 

Klika, J. (1942): Rostlinosociologická studie křivoklátských lesů. Věst. Král. Čes. Společ. Nauk, 
1943, Praha cl. math.-natur. 1941/3: 1–46. 

Marhold, K. & Hindák F. (eds., 1998): Zoznam 
nižších a vyšších rastlín Slovenska. Veda, Bratislava, 687 pp. 

Molnár, A. V. (1999): Bevezetés Magyarország f loristikai növényföldrajzába. In: Sándor, F. (ed.): 
Magyarország védet növényei. Mezőgazda Kiadó, Budapest, pp. 43–52. 

Moravcová-Husová, M. (1964): Die Fagetalia-Gesellschaften des Gebirges Branschauer Wald (Branžovský hvozd) in Wesböhmen. Preslia 36: 272– 

288. 
Moravec J. (1979): Das Violo reichenbachianae-Fagetum – eine neue Buchenwaldassoziation. Phytocoenologia 6: 484–504 

Moravec, J. (ed., 1994): Fytocenologie (nauka o 
vegetaci). Academia, Praha, 403 pp. 

Moravec, J. (2000): Luzulo-Fagetum. In: Moravec, J., 
Husová, M., Chytrý, M. & Neuhäuslová, M. 
(eds.): Hygrofilní, mezofilní a xerofilní opadavé lesy. Academia, Praha, pp. 186–190. 

Moravec, J., Husová, M., Chytrý, M. & Neuhäuslová, 

M. (2000): Hygrofilní, mezofilní a xerofilní 
opadavé lesy. Academia, Praha, 319 pp. 
Mucina, L., Grabherr, G. & Wallnöfer, S. (1993): 
Die Pf lanzengesellschaften Österreichs. Teil 

III. Wälder un Gebüsche. Gustav Fischer Verlag, Jena, 353 pp. 
Tarábek, K. (1980): Klimatickogeografické typy. 
In: Mazúr, E. (ed.): Atlas SSR. SAV a SUGK Bratislava, p. 64. 

ter Braak, C. J. F. & Šmilauer, P. (1998): CANOCO 
Reference Manual and User’s Guide to Canoco 
for Windows. Sotware for Canonical Community Ordination (version 4). Centre of Biometry, 
Wageningen, 353 pp. 

Walnöfer, S., Mucina, L. & Grass, V. (1993): Querco-
Fagetea. In: Mucina, L., Grabherr, G. & Walnöfer, S. (eds.): Die Pf lanzengesellschaften 
Österreichs Teil III Wälder und Gebüsche, Gustav Fischer Verlag, Jena, Stuttgart, New York, 
pp. 8–236. 

Weber, H. E., Moravec, J. & Theurillat, J. P. (2000): 
International Code of Phytosociological Nomenclature. 3rd edition. Journal of Vegetation 
Science 11: 739–768. 

Whittaker, R. H. (1962): Classification of natural 
communities. Bot. Rev. 28: 1–239. 

APPENDIX 

Species in one relevé only: 
E3: Quercus cerris 28 (relevé): 1. 
E2: Acer campestre 16: +, Crataegus sp. 16: +, Euony

mus verrucosus 1: +, Sambucus nigra 3: +, Tilia cordata 

4: 1. 
E1: Brachypodium sylvaticum 3: r, Bromus benekenii 
15: +, Calamagrostis arundinacea 30: r, Crataegus 
laevigata 17: r, C. sp. 16: r, Cystopteris fragilis 11: +, 
Equisetum sylvaticum 27: +, Festuca altissima 11: +, 
Gymnocarpium dryopteris 4: +, Hypericum perforatum 
28: r, Lilium martagon 13: +, Melica nutans 28: +, 
Oxalis acetosella 3: +, Populus tremula 23: +, Pyrus 
communis 17: r, Ribes uva-crispa 14: r, Sambucus 
ebulus 14: +, Senecio nemorensis agg. 31: +, Solidago 
virgaurea 30: r, Stellaria holostea 29: r, S. media 11: +, 
Taraxacum sp. 29: r, Tithymalus cyparissias 15: r, 
Vincetoxicum hirundinaria 15: +, Viola hirsuta 28: r, 
V. mirabilis 19: r, V. sp. 16: r. 
E0: Anomodon attenuatus 24: +, Brachythecium 
populeum 6: +, B. rutabulum 4: +, B. salebrosum 4: 1, 

B. starkei 6: +, Bryoerythrophyllum recurvirostrum 
30: +, Bryum capillare 4: +, B. sp. 11: +, Ceratodon 
purpureus 30: +, Cladonia sp. 4: +, Fissidens bryoides 
4: +, Hedwigia ciliata 1: 1, Homalothecium sericeum 
4: +, Isothecium myosuroides 11: B, Leucodon sciuroides 
1: +, Metzgeria furcata 11: +, Mnium stellare 11: +, 
Plagiochila porelloides 13: +, Plagiomnium laetum 
13: +, P. rostratum 1: +. 
Localities of relevés: 
The header data of relevés are listed in the following order: town or village; locality and habitat; 

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HACQUETIA 3/1 • 2004 

longitude and latitude; age of forest (years); altitude (m); slope (°); aspect (°); relevé area (m2); total cover (%); cover E3 (%); cover E2 (%); cover E1 
(%); cover E0 (%); cover of bare rocks (%); average 
height of tree layer (m); date; author(s) of relevé 
(EB – Eva Belanová, BB – Blažena Benčaťová, RH 

– Richard Hrivnák, KU – Karol Ujházy; in alphabetical order); field number. 
1. 
Hajnáčka village, NNE from the village; Steblová 
skala hill (468 m a. s. l.), SSW from the top of 
the hill, bouldery scree, steep slope under the 
top, copice forest; 19° 58,88257’, 48° 14,78795’; 
50–75; 440; 37; 300; 500; 95; 95; 15; 5; 35; 60; 25; 
11.6.2001; EB, BB, RH, KU; 996. 
2. 
Šurice village, S; Pohanský hrad, E from Soví 
vrch hill, Bouldery scree, slightly convex slope, 
coppice forest; 19° 55,34747’, 48° 12,69303’; 
80–100; 540; 20; 315; 525; 90; 90; 1; 3; 20; 85; 
26–28; 11.6.2001; RH; 997. 
3. 
Šiatorská Bukovinka village, 1,25 km to NE 
from the Karanč hill, commercial forest on 
steep path of northern slope of valley, 20 m 
above the Belina brook; 19° 48,08603’, 48° 
10,05386’; about 70; 370; 29; 335; 450; 100; 98; 
1; 25; 0; 0; 28; 12.6.2001; BB, KU; 18/01 
4. 
Hájnačka village, E; Ragač hill, E, stony-bouldery scree, even slope, fallen stems; -; 90–100; 
430; 23; 20; 450; 80; 80; 10; 20; 15; 60; (15– 
18)23–25; 17.7.2001; RH; 1038. 
5. 
Šiatorská Bukovinka village, SE; Karanč hill, SE, 
end of valley branch under the ridge, stony soil 
disturbed by mouf lons, fallen stems and windfalls; 19° 47,70903’, 48° 9,5993’; -; 590; 21; 350; 
450; 70; 70; 0; 10; 0; 0; 23–25; 12.6.2001; RH; 
1001. 
6. 
Bulhary village, E; Malý Bučeň 
hill, S, narrow 
rocky valley, seed stand; 19° 52,36667’, 48° 
17,99283’; 80–100; 335; 15; 30; 375; 90; 90; 0; 8; 
5; 55; 26–28; 17.7.2001; EB, RH; 1037. 
7. 
Šiatorská Bukovinka village, about 400 m to NE 
from the Karanč hill, commercial forest on the 
path of northern rocky (andesite) slope just 
above the steep valley going to NE from the top 
of Karanč; 19° 47,73895’, 48°9,63647’; 80–100; 
550; 30; 350; 600; 95; 95 ;1; 2; 0; 90; -; 12.6.2001; 
BB, KU; 19/01 
8. 
Šiatorská Bukovinka village, SE; Karanč hill, 
about 800 m NW along the ridge, 70 m from 
state border, slightly cocave even slope; 19° 
47,14375’, 48°9,87477’; (55–65)75–90; 565; 8; 
360; 450; 90; 90; 0; 50; 0; 0; 18–20; 12.6.2001; 
BB, RH; 1003. 
9. 
Obručná settlement, SE; Obručnianska baňa, 
1,35 km to SE from the settlement, commercial 
beech forest on very steep slope just below 
sharp edge of plain, with small basalt boulders 
and stones spread on the soil surface; 19° 
53,48152’, 48° 11,08071’; -; 565; 35; 335; 450; 
85; 80; 0; 35; 0,5; 10; 25; 26.6.2001; KU; 36/01 
10. Obručná settlement, SE; Obručnianska baňa, 
SW from stone-pit, stony or locally bouldery 
scree under the ridge, numerous windfalls; 
19°53,53438’, 48°11,09723’; 60–80(90); 560; 
47; 315; 375; 85; 85; 4; 55; 10; 25; 20–23; 
26.6.2001; RH; 1029. 
11. Šurice village, S; Pohanský hrad, under NW 
margin of plain, moderate slope, locally bouldery scree under the rock wall, coppice forest; 
-; -; 530; 3; 100; 300; 95; 95; 0; 73; 30 (indeterminate); 15; 20; 11.6.2001; BB; 174. 
12. Šurice village, S; Pohanský hrad, under NW 
margin of plain, stony or locally bouldery scree 
under the rock walls, coppice forest; 19° 
55,37868’, 48° 12,37505’; -; 540; 30; 360; 450; 
95; 90; 2; 65; 20; 25–30; 22–24; 11.6.2001; RH; 
998. 
13. Šiatorská Bukovinka village, SE; Karanč hill, 
NE from the top of the hill, even slope under 
the ridge; -; (40)60–95; 700; 25; 45; 450; 80; 
80; 0; 25; 0; 0; 19; 12.6.2001; BB, RH, KU; 
1002. 
14. Bulhary village, E; Veľký Bučeň 
hill, NE, even 
slope under the top; 19° 52,56206’, 48°18,45713’; 
(60)80–90; 475; 15; 360; 400; 85; 80; 1; 80; 1; 0; 
23–24; 17.7.2001; EB, RH; 1033. 
15. Bulhary village, E; Veľký Bučeňhill, NE, slightly 
convex slope under the top; 19° 52,30256’, 
48° 18,41844’; 70–85; 500; 10; 300; 375; 85; 85; 
1; 60; 0; 0; 18–20; 17.7.2001; EB, RH; 1034. 
16. Bulhary village, E; Veľký Bučeň 
hill, NE, near 
cottage, slightly convex even slope; 19° 
52,97132’, 48° 18,62863’; (30–50)60–80; 420; 
2; 135; 400; 90; 90; 1; 80; 1; 0; (18)21–23; 
17.7.2001; EB, RH; 1035. 
17. Hájnačka village, Gortva settlement, E; under 
the saddle between Steblová skala hill and 
Veľká skala hill, shallow valley with transition 
to even slope; -; (30–45)70–90; 400; 15; 270; 
480; 97; 97; 1; 50; 1; 0; 28; 11.6.2001; EB, BB, 
RH, KU; 995. 
18. Večelkov village, W; Medvedia výšina hill, narrow and steep valley, gravelly-stony soil; 19° 
53,39122’, 48° 10,09827’; (60–75)80–100; 605; 
22; 20; 400; 90; 90; 3; 10; 1; 0; 26–28; 26.6.2001; 
RH; 1027. 
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KAROL UJHÁZY, RICHARD HRIVNÁK, EVA BELANOVÁ, BLAŽENA BENČAŤOVÁ: THE BEECH FOREST VEGETATION OF THE CEROVÁ VRCHOVINA MTS. 

19. Večelkov village, W; Medvedia výšina hill, NW 
from the top of the hill, even slope; 19° 
53,27297’, 48° 10,34699’; (60–80)80–100; 595; 
8; 45; 400; 85; 85; 10; 15; 0; 0; 23–26; 26.6.2001; 
RH; 1028. 
20. Šiatorská Bukovinka village, SE; Karanč hill, Ku 
javoru; N from the top of the hill, even, slightly 
undulating slope; 19° 47,81398’, 48° 10,23177’; 
80–100; 460; 20; 360; 400; 98; 98; 0; 2; 0; 0; 30; 
12.6.2001; BB, RH, KU; 1000. 
21. Šiatorská Bukovinka village, about 700 m to 
NW from the Karanč hill, 70 m to the NE from 
the Hungarian boundary, commercial forest 
on the lower part of the even slope with 
frequent andesite stones on the soil surface; 
19° 47,18646’, 48° 9,85181’; about 75; 585; 25; 
355; 400; 90; 90; 0; 7; 0; 15; 23; 12.6.2001; KU; 
22/01 
22. Hájnačka village, Gortva settlement, E; between 
Steblová skala hill and Črep hill, slightly convex soil, rarely stones on the surface; 19° 
59,07076’, 48° 15,12481’; 75–95; 455; 9; 290; 
400; 93; 93; 2; 1; 1; 1; 25–27; 11.6.2001; BB, RH, 
KU; 994. 
23. Hájnačka village, Gortva settlement, E; Steblová 
skala, NE from point of hill, under point 468 m 
a. s. l., even slope; 19° 59,03648’, 48°14,86776’; 
60–90; 430; 20; 60; 400; 95; 95; 0; 2; 1; 0; 25B27; 
EB, BB, RH, KU; 11.6.2001, 993. 
24. Hajnáčka village, NNE; 680 m to the NNE from 
the Steblová skala hill, commercial forest on 
the concave slope on the very beginning of valley with basalt boulders spread on the soil surface; 19° 59,16404’, 48° 15,20051’; -; 375; 25; 
320; 450; 95; 90; 1; 15; 2; 5; 28; 11.6.2001; KU; 
16/01 
25. Nová Bašta, village, W; Dunivá hora hill, NW 
from point of hill, even slope; 19° 54,25343’, 
48° 10,75713’; 70–100; 435; 19; 10; 400; 95; 95; 
2; 25; 0; 0; 30; RH, KU; 26.6.2001; 1031. 

26. Bulhary village, E; Malý Bučeň 
hill, slightly convex steep slope above the bottom of valley; 19° 
52,32978’, 48° 17,59569’; 70–80; 330; 20; 250; 
300; 90; 90; 1; 55; 1; 0; 24–25; EB, RH; 27.7.2001; 
1036. 
27. Šurice village, S; Pohanský hrad hill, N from 
point of hill, beside forest road, even convex 
slope; 19° 55,42239’, 48° 12,76054’; -; 410; 10; 
30; 400; 97; 97; 1; 50; 0; 0; 20; BB; 11.6.2001, 
175. 
28. Nová Bašta, village, W; 750 m to NNE from the 
top of the Medvedia výšina hill, stripe of forest 
between ridge and meadow, moderate convex 
slope with rocky surface; 19° 53,39805’, 48E° 
10,3892’; 40-70; 580; 10; 300; 400; 85; 70; 0,5; 
60; 5; 30; 20; 26.6.2001; KU; 35/01 
29. Šurice village, S; below side ridge of Pohanský 
hrad plain (about 800 m to the west from the 
top of the hill), about 900 m to the north from 
the Garát saddle, commercial forest on steep 
convex slope with shallow rocky red brown soil; 
19° 54,73894’, 48° 12,03533’; about 70; 565; 27; 
315; 450; 95; 90; 0; 30; 10; 35; 20; 11.6.2001; EB, 
KU; 17/01 
30. Nová Bašta, village, W; about 500 m to the E 
from the top of the Medvedia výšina hill, relatively sparse commercial forest on mild and 
convex top part of slope at the edge of ridge 
plain, shallow red brown soil; -; about 80; 625; 
22; 355; 350; 80; 70; 0,5; 30; 5; 5; 20; 26.6.2001; 
KU; 34/01 
31. Obručná settlement, SE; Obručnianska baňa, 
near the side of stone-pit, steep even slope under the ridge, coppice forest; 19° 53,63695’, 
48° 11,15338’; 50–70; 570; 42; 340; 300; 93; 93; 
1; 30; 10; 0; 18; RH, KU; 26.6.2001; 1030. 
Recieved 2. 10. 2003 
Revision recieved 2. 2. 2004 
Accepted 6. 2. 2004 

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