Effect of habitat and historical factors on the distribution of meadow plant species in the radziejowa range (Beskid S Mountains)

Abstract

The main objective of our study was to quantify the effect of habitat factors and methods of land use in the mid-19 century and in the 1980s on the distribution of meadow plants. Based on canonical correspondence analysis (CCA), it was found that altitude, solar radiation and the duration of meadow use had the greatest effect on the species composition of grasslands. Because these factors are often correlated with one another, it is difficult to identify the decisive factor. Many species were limited to areas characterized by long duration of meadow use and unfavourable conditions for agricultural production. For these species to be preserved, an incentive system that gives preference to marginal land is required.

Full text

Introduction
Grasslands, which form a major part of the seminatural
cultural landscape in mountains, are important for preserv-
ing the diversity of plant communities and species. As a
result, agri-environmental programmes that promote grass-
land use have been introduced in many European countries,
including Poland. However, these efforts were often found
to be of low efficiency [1-4], which is probably due to very
general recommendations and the fact that grassland use is
not adjusted to individual communities and species. For this
reason, it is necessary to identify factors affecting grassland
distribution and species composition before detailed pro-
jects are made.
Many authors hold that the main factor affecting species
diversity are soil parameters, in particular the abundance of
nitrogen and phosphorus in the soil, and pH value [5-7]. In
the mountain areas, considerable differences in vegetation
result from the topographic factor [8], i.e. altitude and slope
exposure. The prevalence of plants is also affected by
human activities such as current use (cutting, grazing) [9-
13] and land use history [5, 14].
The objective of the study was to quantify the effect of
different habitat factors and history of past land use on the
prevalence of some meadow plant species in a topographi-
cally diverse area subjected to different utilization methods
in the past.
Material and Methods
The Radziejowa range belongs to the Beskid Sądecki
Mountains, which are part of the Beskid Wysoki Mountains.
Grasslands are found at altitudes ranging from 340 m to
almost 1000 m above sea level. Brown soils proper (leached
and gleyed) occupy the largest area. The mean annual air
temperature ranges from 3ºC on the northern outskirts of
Beskid Sądecki to approx. 1.5ºC in the flat-topped parts of
the mountains. The annual precipitation ranges from 800
mm in Poprad and Dunajec River valleys to 1,100 mm in
the highest parts.
Polish J. of Environ. Stud. Vol. 18, No. 5 (2009), 949-955
Original Research
Effect of Habitat and Historical Factors
on the Distribution of Meadow Plant Species in the
Radziejowa Range (Beskid Sądecki Mountains)
J. Zarzycki1*, M. Kopeć2
1Department of Ecological Bases of Environmental Engineering, Agricultural University in Kraków, Poland
2Department of Agricultural Chemistry, Agricultural University in Kraków, Poland
Received: 10 September 2008
Accepted: 12 January 2009
Abstract
The main objective of our study was to quantify the effect of habitat factors and methods of land use in
the mid-19th century and in the 1980s on the distribution of meadow plants. Based on canonical correspon-
dence analysis (CCA), it was found that altitude, solar radiation and the duration of meadow use had the great-
est effect on the species composition of grasslands. Because these factors are often correlated with one anoth-
er, it is difficult to identify the decisive factor. Many species were limited to areas characterized by long dura-
tion of meadow use and unfavourable conditions for agricultural production. For these species to be preserved,
an incentive system that gives preference to marginal land is required.
Keywords:mountain grasslands, biological diversity, history of land use
*e-mail: janzar@ar.krakow.pl

A total of 350 phytosociological relevés of the
Radziejowa range grasslands were done using the Braun-
Blanquet method [15]. The basic topographic parameters
(altitude, slope and exposure) were determined and a soil
sample was taken from the root zone. These samples were
analyzed for KCl pH and available forms of phosphorus
and potassium using the method of Egner as modified by
Riehm [16]. Solar radiation, which varied according to
exposure and slope, was calculated assuming horizontal
surface solar radiation to be 100%. In the 19th century, land
use method in the study area was based on Austrian cadas-
tral maps with a scale of 1:2880, while data on land use in
the early 1980s were based on topographic maps with a
scale of 1:10,000.
Where a large number of phytosociological relevés
involve tens of species and numerous factors that affect
them, simultaneous analysis of all the relationships is not
possible and in spite of some statistical constraints [17]
multivariate analysis is used [18, 19]. In the present
study, the effect of individual factors on species compo-
sition was analyzed using Canonical Correspondence
Analysis (CCA), which enables the most important gra-
dients in a data set to be identified. Figures show species
that are best adapted to individual environmental factors.
Because nominal variables cannot be used directly in
multidimensional analysis, the use of a point in the past
was noted as dummy variables; they assume 1 if a point
belongs to a certain category, and 0 if it does not. We cre-
ated four variables to illustrate the use of the points in the
past:
GG – grassland in the 19th century and in the 1980s;
AA – arable land in the 19th century and in the 1980s;
GA – grassland in the 19th century and arable land in the
1980s;
AG – arable land in the 19th century and grassland in the
1980s.
Ordination analyses were performed using CANOCO
for Windows ver. 4.5, and the resulting figures were made
using CanoDraw for Windows.
Results
Using the forward selection procedure of CCA analysis
and the Monte Carlo permutation test, exposure and two
variables illustrating past use (AG, GA) were rejected as
they did not significantly increase the percentage of varia-
tion explained. CCA analysis showed that selected habitat
factors account for 13.99% of total variation in species
composition. The effect of particular factors, after eliminat-
ing the effect of others, is shown in Table 1, and the rela-
tionships between factors are illustrated in Fig. 1.
Altitude had the greatest effect on species composition.
The greatest correlation with altitude was shown by bilber-
ry (Vaccinium myrtillus), mat-grass (Nardus stricta) and
broad-leaved meadow-grass (Poa chaixii) in the higher
parts of the massif, and by soft brome (Bromus
hordeaceus), field bindweed (Convolvulus arvensis) and
corn speedwell (Veronica arvensis) in the lower parts of the
massif (Fig. 2). Sickle alfalfa (Medicago falcata), fairy flax
(Linum catharticum) and agrimony (Agrimonia eupatoria)
were found in the areas with higher soil pH, while common
hawkweed (Hieracium lachenalii), mat-grass, heath speed-
well (Veronica officinalis) and oakforest woodrush (Luzula
luzuloides) were associated with higher soil acidity (Fig. 2).
Higher solar radiation values favoured the prevalence of
greater knapweed (Centaurea scabiosa), hoary plantain
(Plantago media) and heath sedge (Carex flacca). True
oxlip (Primula elatior) and spiked rampion (Phyteuma spi-
catum) were found in areas with the lowest solar radiation.
The species associated with high slope inclination were
mountain clover (Trifolium montanum), heath grass
(Danthonia decumbens) and woodland strawberry
(Fragaria vesca). Flat areas were typified by species asso-
ciated with lowland communities such as dandelion
(Taraxacum officinalis), rough bluegrass (Poa trivialis),
950 Zarzycki J., Kopeć M.
Variable Eigenvalues % F P
Altogether 0.388 13.99 5.0 0.01
Altitude 0.09 3.3 11.95 0.01
GG 0.09 3.3 11.15 0.01
AA 0.08 2.9 9.84 0.01
pH 0.07 2.7 9.59 0.01
Insolation 0.07 2.5 9.1 0.01
Inclination 0.04 1.6 5.45 0.01
Soil K 0.03 1.1 3.82 0.01
Soil P 0.02 0.7 2.58 0.01
Table 1. Impact of selected variables on botanical composition.
Eigenvalues – measure for explanatory power of the explanatory
variables (total inertia = 2.725);
% – percentage of explained variance;
F – F ratio for the test of significance of all canonical axes;
P – corresponding probability value obtained by the Monte-
Carlo-permutation test.
-1.0 1.0
-0.6
0.8
Altitude
Inclination
Insolati on
pH
Soil P
Soil K
AA
GG
I ax
IIax
Fig. 1. Ordination diagram of selected variables with axes I.
and II. of a Detrended Correspondence Analysis (DCA).
Environmental factors are represented by arrows. Land use in
the past is represented by stars.

common mouse-ear (Cerastium holosteoides) and bush
vetch (Vicia sepium) (Fig. 3).
The available phosphorus content of soil varied con-
siderably but showed little variation according to species.
Broad-leaved meadow-grass and wild angelica (Angelica
sylvestris) showed the largest correlation with this parame-
ter. Slightly higher species correlations were found for the
available potassium content of soil. The highest correlation
with elevated potassium levels was shown by thermophilic
species such as sickle alfalfa, greater knapweed
(Centaurea scabiosa) and cypress spurge (Euphorbia
cyparissias), and by species such as caraway (Carum
carvi). Species found in areas with the lowest potassium
content were broad-leaved dock (Rumex obtusifolius) and
soft brome (Fig. 4).
Analysis of past land use showed that there were two
distinctive groups of species. First associated with contin-
uous meadow use since the mid-19th century and second
associated with areas that recently served as arable land.
Long-term meadow use is associated to a large extent with
wavy hairgrass (Deschampsia flexuosa), bilberry and
wood betony (Betonica officinalis). The species associated
with the shortest formation of the meadow community
include ruderal species such as broad-leaved dock, broad-
leaved chervil (Chaerophyllum aromaticum) and wild
chervil (Anthriscus sylvestris) (Fig. 5).
Effect of Habitat and Historical Factors... 951
-1.0 1.0
-1.0
1.0
Agr cap
Ant syl
Arr ela
Bro mol
Car aca
Con arv
Cre bieDac glo
Dau car
Her sph
Hol lan
Hol mol
Luz luz
Lyc lcu
Nar str
Poa cha
Pot ere
Tar o ff
Tri dub Tr i pr a
Vac my r
Ve r a rv
Altitude
-1.0 1.0
-1.0
1.0
Car fla
Car car
Cre bie
Cyn cri
Dac glo
Fes pra
Gal mol
Hie lach
Hol mol
Hyp mac
Lat prat
Luz luz
Nar str
Pim maj
Pla med
Pot ere
Ran pol
Rum ace
Tar of f
Vac myr
Ver of f
pH
AB
Fig. 2. Ordination diagram (CCA) of best-fitted species to altitude (A) and pH (B).
-1.
0
1.
0
-1.0
1.0
Ast maj
Bri med
Car aca
Cre mol
Dau car
Gal mol
Gen asc
Hie la ch
Hyp ma c
Kna arv
Leo his
Lot co r
Luz luz
Phy spi
Pim sax
Pla med
Pri ela
Ran acr
Ran pol
Thy pul Tri mon
Insolation
-1.
0
1.0
-1.0
1.0
Bri med
Cer hol
Cre bie
Dan dec
Fra ves
Leu vul
Luz luz
Lyc lcu
Phl pra
Pim sax
Poa tri
Pot ere
Ran rep
Rhin m in
Ta r of f
Thy pul
Tri mon
Tri rep
Tris fla
Vic se p
Inclination
AB
Fig. 3. Ordination diagram (CCA) of best-fitted species to insolation (A) and inclination (B).

Discussion of Results
Because the analyzed factors are often strongly correlat-
ed, it is not possible to identify the dominant factor because
the species composition of plant communities is the result of
habitat factors and method of use [3]. There was no available
information about present management of grasslands (e.g.
level of fertilization, number of cuts etc.), which is an impor-
tant factor determining the species composition. In effect, the
percent of variation explained is relatively low. Based on the
present results, two basic groups of species related to the first
DCA axis can be identified, which correspond to the course
of largest species variation. The first group includes species
found in the highest parts of the range, on highly inclined
slopes with strongly acidified soils. These conditions also
involve the long-term use of land as grasslands. Similar rela-
tionships between the duration of meadow use and altitude
and soil pH were obtained in Norway [5] and in Germany
[13]. Such areas comprise species typical of sparse mat-grass
swards (Nardo-Calunetea): mat-grass and tormentil
(Potentilla erecta); and spruce forests (Vaccinio-Picetea):
bilberry and wavy hairgrass.
952 Zarzycki J., Kopeć M.
-1.0 1.0
-1.0
1.0
Ang syl
Arr ela
Bri med
Cam per
Car hal
Car pal
Cyn cri
Hyp mac
Leo his
Poa cha
Rum ace
Ver ch a
P
-1.
0
1.
0
-1.0
1.0
Agr eup
Bri med
Bro mol
Car hal
Car fla
Car aca
Car car
Cen sca
Cir arv
Eup cyp
Gal mol
Hyp rad
Kna arv
Lin cat
Lyc lcu
Med fal
Phl pra
Pla med
Ran pol
Ran rep
Rum ace
Rum obt
Thy pul
Vio can
K
AB
Fig. 4. Ordination diagram (CCA) of best-fitted species to available soil P content (A) and K content (B).
0.0 1.0
-1.0
0.4
Ant syl
Arr ela
Cam pat
Cer hol
Cha aro
Cre bie
Dac glo
Dau car
Her sph
Hollan
Hyp rad
Phl pra
Poa tri
Ran rep
Rum obt
Tar of f
Tri dub
Tri pra
Tri rep
Vic se p
AA
0.0 1.0
-0.4
1.0
Agr cap
Ast maj
Bet off
Car pil
Cen jac
Cre mol
Cru gla
Dan dec
Des fle
Gen asc
Hie lach
Hie pil
Luz luz
Nar str
Poa cha
Pot ere
Vac my r
Ver off
GG
AB
Fig. 5. Ordination diagram (CCA) of best-fitted species to short grassland use (A) and long grassland use (B).

Lower altitudes are dominated by habitats with low
inclination and higher soil pH. These habitats are charac-
terized by relatively good productive properties, which is
why they were recently used as arable land. They were
changed into grassland by sowing meadow mixtures or
through spontaneous revegetation. In most cases, these are
unstabilized communities. The species found under these
conditions include plants typical of lower communities of
the order Arrhenatheretalia: cocksfoot (Dactylis glomera-
ta), meadow fescue (Festuca pratensis), tall oat-grass
(Arrhenatherum elatius), and Queen Anne’s lace (Daucus
carota). Ruderal species such as field bindweed, broad-
leaved dock and wild chervil are also found. Differences in
habitat fertility also vary according to the main gradient of
plant variation. In many cases, this factor is considered
decisive for the occurrence of meadow species [6, 20].
The second DCA axis is correlated with the content of
available phosphorus and potassium and solar radiation.
Only the group of thermophilic species occurs on soils with
a higher potassium content. These include agrimony, sickle
alfalfa and dodder (Cuscuta epithymum), which are com-
mon on the meadows of the neighbouring Pieniny
Mountains. Unlike the results of many meadow vegetation
studies in Western Europe, where high phosphorus content
is considered an important factor reducing species diversi-
ty [20-23], influence of this factor on botanical composition
of grassland sward in Radziejowa Range is low. One possi-
ble explanation is the presence of phosphorus-deficient
soils in the Beskid Sądecki and the low level of phosphorus
fertilization. It is in accordance with results obtained by
many authors [11, 20, 23, 24]. They found a significant
effect of available phosphorus content on botanical compo-
sition of grassland but only in the case of high phosphorus
content. The analysis revealed a few species connected with
the naturally most interesting mountain meadow communi-
ty (Gladiolo-Agrostietum). The main reason is that they
Effect of Habitat and Historical Factors... 953
Agi eup Agimonia eupatoria Dan dec Dantonia decumbens Phy spi Phyteuma spicatum
Ang syl Angelica sylvestris Dau car Daucus carota Pim maj Pimpinella major
Ant syl Anthriscus sylvestris Des fle Deschampsia flexuosa Pim sax Pimpinella saxifraga
Arr ela Arrhenatherum elatius Eup cyp Eupatoria cyparisias Pla med Plantago media
Ast maj Astrantia major Fes pra Festuca pratensis Poa cha Poa chaixi
Bet off Betonica officinalis Fra ves Fragaria vesca Poa tri Poa trivialis
Bri med. Briza media Gal mol Galium mollugo Pot ere Potentilla erecta
Bro mol Bromus mollis Gen asc Gentiana asclepiadea Pri ela Primula elatior
Cam per Campanula persicifolia Her sph Heracleum sphondylium Ran rep Ranuculs repens
Car hal Cardaminopsis halleri Hie lac Hieracium lachenali Ran acr Ranuculus acris
Car fla Carex flava Hie pil Hieracium pilosella Ran pol Ranunculus polyanthemos
Car pil Carex pilulifera Hol lan Holcus lanatus Rhi min Rhinanthus minor
Car aca Carlina acaulis Hol mol Holcus mollis Rum ace Rumex acetosa
Car car Carum carvi Hyp mac Hypericum maculatum Rum obt Rumex obtusifolius
Cen jac Centaurea jacea Hyp rad Hypochoeris radicata Thy pul Thymus pulegiodes
Cen sca Centaurea scabiosa Kna arv Knautia arvensis Tri mon Trifolim montanum
Cer hol Cerastium holosteoides Lat pra Lathyrus pratensis Tri dub Trifolium dubium
Cha aro Chaerophyllum aromaticum Leu vul Leucanthemum vulgare Tri pra Trifolium pratense
Cir arv Cirsium arvense Lin cat Linum catharticum Tri fla Trisetum flavescens
Con arv Convolvulus arvensis Lot cor Lotus corniculatus Vac myr Vaccinium myrtillus
Cre bie Crepis bienis Luz luz Luzula luzuloides Ver arv Veronica arvensis
Cre mol Crepis mollis Lyc lcu Lychnis flos-cuculi Ver cha Veronica chamaedrys
Cru gla Cruciata glabra Med fal Medicago falcata Ver off Veronica officinalis
Cyn cri Cynosurus cristatus Nar str Nardus stricta Vic sep Vicia sepium
Dac glo Dactylis glomerata Phl pra Phleum pratense Vio can Viola canina
Table 2. Abbreviations of species names used in Figures.

occur when the analyzed factors were at a medium level.
Only brown knapweed (Centaurea jacea), colonial bent-
grass (Agrostis capillaris) and Northern Hawk’s-beard
(Crepis mollis) were associated with long-term meadow
use.
Conclusions
A large number of meadow species are only found in
communities that are marginal from the agricultural point
of view (high altitude, low pH, steep slope) and have long
been used as meadows and pastures. A similar relationship
that the high nature quality species are found mainly in
undisturbed natural or semi-natural habitats has been
reported by other authors [24, 25]. The socio-economic
processes result in such areas being abandoned or afforest-
ed, which takes place in Poland and other European coun-
tries [26-29]. Feed production in the Beskidy Mountains is
now carried out mainly on old arable land characterized by
better production conditions, but the plants that grow there
usually represent common species. For many species that
have adapted to specific habitat conditions and show low
ecological amplitude to be preserved, it is necessary to use
a system of payments that gives preference to the use of
meadows and pastures at higher altitudes and on poorer
soils, even at the cost of reducing subsidies on grasslands
on old arable land of low natural value. Result-orientated
subsidies in grasslands should be created and the farmers
are to be paid for keeping up a high plant species richness
[4, 13]. In case of well prepared, prescriptions the results
can be satisfactory [30].
Acknowledgements
This article was prepared as a part of research project
No. 2P06S 01228 financed by the State Committee for
Scientific Research.
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