journal 26.indb

POLISH JOURNAL OF ECOLOGY
(Pol. J. Ecol.)
59
2
249–262
2011
Regular research paper
Alina STACHURSKA-SWAKOŃ* and Krystyna KUŹ
Institute of Botany, Jagiellonian University, Kopernika 27, 31–501 Kraków, Poland,
*e-mail: [email protected] (corresponding author)
PHENOTYPIC RESPONSE OF DORONICUM AUSTRIACUM JACQ.
(ASTERACEAE) TO DIVERSE MOUNTAIN AND LOWLAND
CONDITIONS
ABSTRACT: The knowledge of phenotypic
response of rare and protected species provide
useful information for the conservation and
management strategies. Doronicum austriacum,
a subalpine Central-European species has several
lowland localities, which in Poland are regarded
as glacial relicts. Diverse edaphic, climatic and
coenotic conditions in particular localities give
evidence to the broad ecological amplitude of the
species. Based on data pertaining to three various
populations occurring in different geographical
regions (from South and Central Poland), different elevations a.s.l. (275–1350 m a.s.l.) and growing within different plant communities (subalpine
tall-herb communities, mountain meadow, carr)
an attempt has been made to characterise selected
morphological, developmental and ecological features (like number of capitula and their diameter,
effectiveness of reproduction, spatial distribution)
and to test a hypothesis as to whether a lowland
form of this species exists. The results indicate certain statistically significant differences (as number
and diameter of capitula) between the study populations pertinent to plants at the generative stage.
The distribution of the examined traits, however,
falls within the range of species variability. The
differences between averages are conditioned by
the quality of the environment in terms of the soil
moisture level, availability of mineral compounds
and lighting, as well as by the degree of competition from other plants. The result of the experiment, involving transplantation of specimens into
journal 26.indb 249
gardens, allow for the presumption that morphological features undergo environment-related
modifications. At this stage of the studies, the idea
of a morphologically different lowland form of the
species cannot be supported.
KEY WORDS: Doronicum austriacum, subalpine species, isolated population, biometry, ecological amplitude, plasticity, macroforbes, Carpathians
1. INTRODUCTION
The phenotypic plasticity is a fundamental property of plant species (S ch li cht i ng
1986, 2002, Mi ne r et al. 2005, Pi g l iuc c i
2005, Va l l a d are s et al. 2007). The morphological traits of plants are affected both by
abiotic factors like light, temperature, water
and nutrients availability as well as biotic factors including the role of competitors, predators and pollinators. The variability of traits
along the environmental gradient is particularly interesting in plants with isolated populations when the site conditions are extremely
different from range of the species. In case of
rare or protected species the knowledge of
their phenotypic response has the potential
to provide useful biological information for
the conservation and management strategies
2011-06-28 10:20:20
250
Alina Stachurska-Swakoń and Krystyna Kuź
(Mit k a and Tu m i d aj ow i c z 1993, A l e r i c
and Ki rk man 2005, Hens en and Wes che 2006, Kost ra k ie w ic z 2009).
Doronicum austriacum Jacq., Austrian
leopard’s bane, is one of many Central-European species with the centre of its distribution
in the mountains of Europe: the Carpathians,
Balkans, Alps, Apennines and the Eastern Pyrenees (Kucowa 1971, Á lvare z Fer nánd e z
2003). In Poland, the species occurs mainly
in the Carpathians and rarely in the Sudety
Mts., where it occurs only in central and eastern ranges (Z aj ąc and Z aj ąc 2001). However, there are some isolated populations of
the species situated outside of the continuous
range in the mountains. The localities north
of the arch of the Carpathians are located in
the mountain regions in Central Poland (the
Świętokrzyskie Mts.) and in Southern Poland
(the Silesian Upland, the Błędów Basin in the
Kraków-Częstochowa Upland). The localities
outside the Carpathians are regarded as glacial relict type localities, and the populations
occurring there – as populations originating
from the Carpathians (Szafer 1930, 1977).
Inventories of lowland localities drawn up in
the early 1990s showed a dramatic decrease in
both the number of existing populations and
the number of individuals of the species in
particular localities; they also indicated real
and potential threats (Bróż and Pr z e my s k i
1992).
Austrian leopard’s bane is regarded as
a characteristic species of macroforb communities of the Adenostylion alliariae Br.-Bl.
1925 alliance (Me dwe cka-Kor naś et al.
1977, Matuszk ie w icz 2005, St achu rsk a Swa koń 2009). In the lower montane forest
zone in the Carpathians, it co-participates
in forming the Arunco-Doronicetum austriaci association (Kor naś and Me dwe ckaKor naś 1967). In lower mountain sites, it
grows sporadically in moist sycamore-beech
forests, whilst in the Bieszczady Mts. (SouthEast Poland), it is also found in alder woods
representing the associations of Caltho laetae-Alnetum (Zarz. 1963) Stuchlik 1968
and Alnetum incanae Lüdi 1921 (Jasie w i c z
1965, Mi cha li k and Szar y 1998). In isolated relict localities, it grows in cool, swampy
river valleys, mainly in the Fraxino-Alnetum
W.Mat. 1952 association (Bróż and Prz emysk i 1992). The locality in the former
journal 26.indb 250
spring area of the Biała river in the Błędowska
Desert is an exception. The locality was discovered in 1924 by Pi e ch (1924). The author wrote about ‘abundant forest vegetation’,
which had developed due to an ’abundance
of water in sandy soil’. He listed Alnus glutinosa (L.) Gaerth., Corylus avellana L., Frangula alnus Mill., Ribes sp., and Salix sp. div.
among the plants of the stream terrace and
mentioned the presence of ‘humus typical of
moist forests’. In 1975, the springs of the Biała
river and its tributaries disappeared because
of the lowering ground-water table following
the development of industrial plants and the
opening of the mine (“Pomorzany”, Wój ci k
1997). At present, forest communities are
greatly depleted, and the area occupied by
the population of D. austriacum has been invaded by coniferous forest species and Calamagrostis epigejos (L.) Roth. Nevertheless, the
population of Doronicum still remains there
despite dramatic changes in substrate moisture and the quality of the communities. The
above data indicate a wide broad ecological
scale of the species and its ability to survive in
changing environmental conditions.
In the study on the springs of the Biała
river, Pi e ch (1924) noted morphological differences between the specimens examined
there and the plants growing in the mountains. The differences in the extrazonal appearance of specimens in the Świętokrzyskie
Mts. and the Silesian upland were also noted
by Bróż and Prz emy s ki (1992), who even
questioned the taxonomical identity of the
species.
The objectives of these studies are: 1) to
determine the scope of selected morphological and developmental features in the relict
population versus mountain populations;
2) to provide ecological characteristics of the
studied populations, and 3) to verify the hypothesis that there is a separate lowland form
of the species.
2. THE STUDY SPECIES
Doronicum austriacum Jacq. (Asteraceae) has a short rhizome without runners
with numerous adventitious roots. The
abundantly leaved stem grows up to 120 cm
high (Kucowa 1971). In the general distribution range, some reports say even of the
2011-06-28 10:20:20
Phenotypic reaction of Doronicum austriacum
height above 150 cm (Á lv are z Fer nánde z
2003). In the top part, the stem branches into
a sub-umbelliferous form. The leaves have
variable shapes. The basal leaves are ovate,
obtuse, with shovel-cordate base and blunt
apex, and petiolate. The lower cauline leaves
can be similar to the basal leaves or sessile.
The lower and middle cauline leaves have
fiddle-shaped blades, they are semi-amplexicaul and their dimensions are 6.5–19.0 cm
× 4.5–12.5 cm. The upper cauline leaves are
small: 2.5–13.0 cm × 0.7–5.0 cm, they are
ovate with an acute apex and amplexicaul.
The capitula, 5–7 cm in diameter, sit on long
peduncles. The female flowers are yellow, and
ligulate, and the bisexual flowers are disc florets. The fruit is a single-seed, ribbed achene,
up to 2 mm long. D. austriacum produces two
types of achenes: those developing from disc
florets have pappus and stiff hairs on the ribs
whereas those developing from ligulate florets are naked and without pappi.
3. STUDY AREA
Three populations were selected for this
study (Fig. 1):
1. Lowland population deemed to
be a relict, situated in the former
spring area of the Biała river on the
Błędowska Desert (N50°20’, E19°31’).
The site is situated at 275 m a.s.l.
251
Doronicum austriacum grows within
the former spring area and the terraces of the uppermost sections of
the stream course. The location is
overshadowed mainly by alder trees
as well as hazel and alder buckthorn
shrubs. The plant community on
the site was a degraded riverine carr.
In this paper it is designated as the
BIAŁA population;
2. Mountain population on the Polana
Bieniowa glade in the Gorce Mts.
(N49°33’, E20°10’). The site is situated
in the lower montane forest zone at
1100 m a.s.l. Here the species grows
abundantly in a fully sun-illuminated
site, within the macroforb association
Arunco-Doronicetum austriaci Kornaś
(1955 n.n.) 1967. This site is designated as the GORCE population;
3. Mountain population in the Babia
Góra Mt. massif (N49°34’, E19°31’).
The location is situated in the upper
montane forest zone at 1350 m a.s.l.
Here, Doronicum grows on a moist
slope partly overshadowed by spruces, within the macroforb association Ranunculo platanifolii-Adenostyletum alliariae (Kr aj i na 1933)
Dúbravcová et Hadač ex Kočí 2001.
This site is designated as the BABIA
GÓRA population.
Fig. 1. Locations of the study populations of Doronicum austriacum. The gray color indicates the distribution of the species in Poland.
journal 26.indb 251
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Alina Stachurska-Swakoń and Krystyna Kuź
4. MATERIAL AND METHODS
4.1. Ecological and population data
To estimate the habitat conditions, soil
analyses were completed for each of the localities following Kow a l kow sk i et al. (1973),
L it y ńsk i et al. (1976), Oleksy nowa et al.
(1976). The analyses included, inter alia: organic matter content, organic nitrogen, and
soil absorbing complex. These studies were
conducted in 2006–2007.
During the flowering period, 50 generative shoots and 50 vegetative shoots of Doronicum austriacum were randomly selected
on each location. The height and number of
leaves were determined for each shoot along
with the dimensions (length and width) of
the lowest leaf and one middle leaf (always
the fifth from the bottom). Basal leaves wither fairly rapidly, so during the flowering period they have already withered, therefore their
variability was not taken into account. In
generative shoots, the number of capitula was
counted and their diameters measured. All
measures were taken on living plants growing in the wild. In each location, 5 small 1 m2
plots were selected, again randomly, where
the shoots in a generative or vegetative stage
were counted. The biometric studies were
carried out during season 2006 and 2007 in
BIAŁA site and in season 2007 in BABIA
GÓRA and GORCE sites. The effectiveness
of reproduction was also estimated through
measuring the germinating power. For this
purpose, the collected seeds were sown either
without freezing or after a two-day freezing
period at –15°C in three repetitions, at different intervals after their collection. The seeds
from the GORCE Mts. site were sown after
two, three and four months following collection (with 35 seeds in each portion), whereas
the seeds from the BABIA GÓRA population
were sown after one, three and six months
following collection (with 25 seeds in each
portion). Owing to the small number of seeds
collected from the BIAŁA population, these
were not included in the tests for germinating power. The seed germination experiment
was conducted in the laboratory on moist filter paper lined in 7-cm diameter petri-dishes.
Only several individual plants were observed
after they had been transplanted from the
journal 26.indb 252
BIAŁA site to a substitutive habitat (home
gardens).
The biology of this species is not well
known, so the studies did not include the features related to age, only the shoots were separated into vegetative and generative categories.
4.2. Statistical analysis
The results were submitted to statistical
analyses using the MVSP (Kovach 2007)
and Statistica packages. The quantitative data
showed normal distributions. Apart from basic descriptive statistics (average, maximum,
minimum, standard deviation, and the coefficient of variation = CV%), the data were
also submitted to analysis of variation by oneway ANOVA, in order to test the hypothesis
regarding the lack of differences between the
populations under study. The statistical significance of yearly differences in the BIAŁA
site, where the studies were conducted during two seasons, was checked using a posteriori LSD (Least Significant Difference) test.
The mutual relationships between shoots was
analysed by the dentrended canonical analysis – DCA method.
5. RESULTS
5.1. Soil properties
The basic soil analyses indicated evident
differences in soil quality at the study sites.
The soil from the BABIA GÓRA site was the
richest in terms of mineral and organic components (Table 1). The organic carbon content was 30.4%, nitrogen – 1.7%, and that of
organic matter, 52.2% and it also contained
large amounts of available substances such as
K2O and P2O5. The lowest contents of organic
compounds were noted in the soil in BIAŁA
site. There, only the content of CaO and the
pH level were at their highest values at 103.6
mg CaO, and pH KCl 5.06, respectively. The
hydrolytic acidity of the soil is associated
with the presence of hydrogen and aluminium ions in the absorbing complex of the soil.
It is closely related to the total sulphur content and assumes the values inversely proportional to the pH values. Therefore, the highest
acidity of 61.86 mg occurred in the soil from
BABIA GÓRA site that also had the lowest
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Phenotypic reaction of Doronicum austriacum
253
Table 1. Soil properties of the study sites of Doronicum austriacum.
Org.
matter
(%)
C org.
Sites
Biała
3
1.6
0.03
Gorce
9
5.4
Babia Góra
52
30.4
Available substances (mg·100 g-1)
N org.
(g·100 g-1)
C/N
pH
K 2O
P2O5
CaO
H2 O
KCl
53
0.2
4.6
103.6
5.71
5.06
0.40
13
6.0
2.4
14.0
4.27
3.65
1.69
18
32.2
17.6
33.6
4.05
3.34
Table 2. Absorbing complex of soil in the study localities of Doronicum austriacum. H – hydrolytic acidity, S – total of ion-exchange bases, T – exchange capacity, V – level of soil saturation with bases.
Sites
H
Soil absorbing complex (mg·100 g-1)
S
T
Biała
2.38
9.00
11.38
79
Gorce
16.89
4.10
20.99
20
Babia Góra
61.86
29.64
91.50
32
pH, whereas the lowest acidity (2.38 mg) occurred in the soil from BIAŁA site that has
the highest pH. The highest total of ion-exchange bases – 29.64 mg was found in the soil
from BABIA GÓRA site, the lowest (4.10 mg)
– in the soil from the GORCE site.
The measure showing the capacity to absorb ions is the exchange capacity expressed
in mg per 100 g of soil. The percentage ratio
of the total amount of basic cations absorbed
in the soil to the exchange capacity (T) is an
indicator of the level of soil saturation with
bases (V). The exchange capacity of sands
is 1–10 – as in the case of the soil from the
BIAŁA site, loam soils – 10–20 (GORCE
site) clay soils – up to 60, and peats – even
150 and more. As a rule, the degree of acidic soil saturation with bases in pine forests
is low (such as in the case of BABIA GÓRA
site – 32.39 and GORCE site – 19.53 mg), and
much higher in the neutral or alkaline soils
of deciduous woods – namely in BIAŁA sites
where it stands at 79.09 mg (Table 2). The degree of saturation of the absorbing complex
by basic cations determines the buffer capacity of the soil.
5.2. Morphological features
The heights of the generative shoots of
Doronicum austriacum under study ranged
from 51 (GORCE site) to 120 cm (BIAŁA
journal 26.indb 253
V(%)
site). On average, the tallest generative
shoots were recorded in BIAŁA population –
90.4 cm (CV = 79%), and the shortest – in the
GORCE population – 72.3 cm (CV = 19%;
Fig. 2A). The vegetative shoots were markedly shorter, ranging from 29 cm (GORCE
population) to 97 cm (BABIA GÓRA population). The lowest average for vegetative shoots
was recorded in BIAŁA population – 48.2 cm
(CV = 20%), whereas the highest was at BABIA GÓRA population – 56.6 cm, where the
dispersion of values describing this feature
was also the broadest (CV = 31%, SD = 14.2;
Fig. 2A). The number of leaves on generative
shoots ranged from 5 (BIAŁA population)
to 20 (GORCE population). On average, the
highest number values of leaves were found
on generative shoots in GORCE population
– 13.3, whereas the lowest ones were found in
BIAŁA population – 10.9 (Fig. 2B). The vegetative shoots had from 5 (BABIA GÓRA and
GORCE populations) to 15 leaves (BABIA
GÓRA and GORCE populations). The highest average number of leaves was found in
the BABIA GÓRA population – 10.3, and the
lowest in GORCE population – 9.6 (Fig. 2B).
The length of the lowest leaves on generative shoots fluctuated within the 7.5–20.0 cm
range (from GORCE to BABIA GÓRA population, respectively). The average values were,
however, close to the values found in BIAŁA
population and BABIA GÓRA sites amount-
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Alina Stachurska-Swakoń and Krystyna Kuź
Fig. 2. Biometric traits of Doronicum austriacum shoots in the studied populations.
A – height of shoots, B – number of leaves, C – length of the lower leaf, D – width of the lower leaf,
E – length of the middle leaf, F – width of the middle leaf, G – number of inflorescences, H – diameter
of the capitula.
Bg – BABIA GÓRA population, generative shoots, Bv – BABIA GÓRA population, vegetative shoots,
Gg – GORCE population, generative shoots, Gv – GORCE population, vegetative shoots, Pg1 – BIAŁA
population in the season 2006, generative shoots, Pg2 – BIAŁA population in the season 2007, generative shoots, Pv1 – BIAŁA population in the season 2006, vegetative shoots, Pg2 – BIAŁA population in
the season 2007, vegetative shoots.
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Phenotypic reaction of Doronicum austriacum
255
Table 3. Interset correlation of some variables with DCA axes.
Variable
Height
Number of leaves
Lenght of lower cauline leaf
Width of lower cauline leaf
Lenght of middle cauline leaf
Width of middle cauline leaf
Diameter of the capitula
Number of capitula per one shoot
Axis 1
0.064
0.152
0.000
0.003
0.086
0.077
0.231
0.772
Axis 2
0.000
0.076
0.132
0.166
0.217
0.179
0.111
0.097
Axis 3
0.051
0.059
0.089
0.000
0.159
0.182
0.142
0.183
Table 4. One-way ANOVA results for variables of Doronicum austriacum (P <0.001).
Variable
Height
Number of leaves
Lenght of lower cauline leaf
Width of lower cauline leaf
Lenght of middle cauline leaf
Width of middle cauline leaf
Diameter of the capitula
Number of capitula per one shoot
SS
11284
139.42
334.22
73.08
108.37
53.85
97.85
773.25
df
3
3
3
3
3
3
3
3
MS
3761
46.47
111.41
24.36
36.12
17.95
32.61
257.75
F
17.54
9.49
18.0
10.3
7.49
8.49
48.07
63.29
R2
0.999
0.997
0.999
0.998
0.985
0.993
1
1
Fig. 3. DCA diagram of Doronicum austriacum generative shoots along 1st and 2nd axis. Circles – BABIA
GÓRA population, squares – GORCE population, triangles – BIAŁA population.
ing to 13.9 cm and 13.5 cm, respectively, but
in the GORCE population, the average value
is much lower at 10.7 cm (Fig. 2C). The width
of the lower leaves fell within the 4.0–13.4 cm
range (GORCE and BIAŁA population, respectively). The average value was the highest
journal 26.indb 255
at BABIA GÓRA population – 8.1 cm, and
the lowest in GORCE population – 6.4 cm
(Fig. 2D).
The length of the middle leaf ranged from
6.5 cm in GORCE population to 21 cm at BABIA GÓRA population. The lowest average
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Alina Stachurska-Swakoń and Krystyna Kuź
der study, with more similarity between the
shoots from BIAŁA population and BABIA
GÓRA sites than between BABIA GÓRA
and GORCE sites. The first and second axes
explain 52% and 17% of observed variability, respectively. The most significant are
the generative features, and in particular the
number of inflorescences. The effect of the
study features on grouping is shown in Table
3. There is no grouping whatsoever of vegetative shoots (Fig. 4).
On the basis of the selected features, the
results of ANOVA do indicate statistically
significant differences between the studied
populations (Table 4). The significance of differences between years was also tested, but
these turned out to be statistically insignificant. The significance of the differences could
only be tested for the BIAŁA population, because it was the only one where the studies
were carried out in two seasons.
length of the middle leaf equal to 11.6 cm was
found in BIAŁA population, whereas the highest one close to 13.6 cm was found at BABIA
GÓRA site (Fig. 2E). The width of the leaf correlated with its length, the average being 3.6 cm
in the BIAŁA population and 7.9 cm in the BABIA GÓRA population (Fig. 2F). The length of
middle leaves on the vegetative shoots fell within the 6–15 cm range (BIAŁA population and
BABIA GÓRA, respectively). The lowest average was found in the leaves from BIAŁA population, and the highest – from BABIA GÓRA
population. The width of this leaf was highest in
GORCE site, and lowest in BIAŁA population
(Fig. 2F). The width values showed less variability than leaf length values.
The largest differences were seen in the
numbers of inflorescences. The number of
capitula per one generative shoot ranged from
1 to 16. The maximum number was found in
GORCE population (16) as well as the mean
value (7). It was interesting to find that there,
in GORCE site, were no shoots with less than
three capitula. The lowest number of inflorescences on a single shoot was found in
BIAŁA population – 2.3 (with the 1–7 range;
Fig. 2G). The diameter of the capitula ranged
from 2.0 to 7.7 cm, with the smallest capitula developing in the BIAŁA population was
equal to 3.7 cm, and the largest in GORCE
population – 5.4 cm (Fig. 2H).
The diagram of DCA analysis was prepared on the basis of features found for
generative shoot groups of particular populations (Fig. 3). However, there are no clearcut differences between the populations un-
5.3. Group features
The populations under study showed
clustered spatial distribution, which was most
notable in the GORCE population. There, the
population was also the most numerous. On
the BABIA GÓRA site, the individuals of Doronicum austriacum grew either singly or in
smaller clusters.
The number of generative shoots was
higher than the vegetative ones in the GORCE
and BABIA GÓRA sites. On the BIAŁA population, there was a prevalence of vegetative
shoots (Table 5).
Table 5. Number (shoots m-2 ) of vegetative and generative shoots of Doronicum austriacum on five plots
(1 m2) in each study sites.
sites
Shoots
Plots
Mean
1
2
3
4
5
vegetative
7
7
8
10
9
8.2
generative
4
9
3
7
6
5.8
vegetative
8
4
2
3
3
4.0
generative
9
5
11
12
11
9.6
vegetative
4
3
2
2
4
3.0
generative
3
3
5
3
6
4.0
Biała
Gorce
Babia Góra
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2011-06-28 10:20:22
Phenotypic reaction of Doronicum austriacum
257
5.4. Reproduction effectiveness
5.5. Ex-situ plant development
In the pilot studies concerning reproduction, seeds were collected from the GORCE
and BABIA GÓRA populations. Owing to the
shortened vegetation period (drought), the
seeds from the BIAŁA population could not
be collected in sufficient numbers. In general, the capitula from the GORCE site had
the highest (estimated) numbers of achenes.
These achenes were also larger and thicker
than the seeds collected from BABIA GÓRA
site (also only estimated). These achenes,
however, often contained seed-eating insect
larvae. The seeds from the BIAŁA population
site were the smallest, in relative terms.
In the samples, both frozen and nonfrozen seeds germinated. The first seedlings
emerged ca 8 days after sowing, and the germination lasted ca 7 days. The seeds from
GORCE site germinated faster and there was
also a higher percentage of germinating seeds.
The germinating power after freezing was
higher in the case of the seeds from BABIA
GÓRA site, whereas among the seeds from
GORCE site, it was higher in the group that
was not frozen (Fig. 5). Also, these preliminary results might indicate that the viability
of seeds decreases with time.
These observations were conducted in
two home gardens: in Wojnicz (the Wielickie
Foreland, Western Carpathians) and Andrychów (the Beskid Mały Mts., Western Carpathians), on several individuals (5 in each
garden) transferred from the BIAŁA population at the beginning of vegetation season
in 2006. These individuals had already developed basal leaves. The growth started in
April and proceeded rapidly. The beginning
of flowering was noted as early as towards
the middle of May and lasted till late June. In
2006, all the individuals had 3–5 capitula. The
number of capitula increased in subsequent
years: to 4–8 in 2007, and from 4 to 10 in
2008. The leopard’s bane cultivated in garden
conditions (regular watering and fertile soil)
reaches 65 to 120 cm in height. The average
number of leaves did not change significantly
in the subsequent years of observation, remaining at from 13–17.
6. DISCUSSION
The geographical distance between the
BIAŁA population and the continuous distribution range in Carpathians as well as iso-
Fig. 4. DCA diagram of Doronicum austriacum vegetative shoots along 1st and 2nd axis. Circles – BABIA
GÓRA population, squares – GORCE population, triangles – BIAŁA population.
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Alina Stachurska-Swakoń and Krystyna Kuź
Fig. 5. Percent of frozen and un-frozen seed germination of Doronicum austriacum from BABIA GÓRA
and GORCE populations in different time of sowing after collection.
lated populations in the Świętokrzyskie Mts.
allows one to assume that there is a barrier
preventing gene flow between these populations. Therefore, the BIAŁA population
can be considered to be a closed population
(May r 1974), where genetic phenomena associated with small populations occur, such
as: the founder effect, genetic drift, inbreeding, or an increase in homozygosis (Mit ka
1997, A l l endor f and Lui kar t 2007). More
detailed studies will be needed to find whether speciation processes are currently at work
in this population.
The results presented in this study indicate the wide ecological scale of the species
concerned. The sites selected for the study
differed significantly with respect to habitat
conditions: climatic (elevation a.s.l. and related parameters: temperature, amount of precipitation, duration of vegetation season, etc.)
and microclimatic (influx of solar radiation
resulting from shadow or its lack of), soil and
coenotic conditions. On the basis of the examined features, the results observed indicate
statistically significant differences between
the study populations. The differences are
evident for generative shoots, whereas they
are vague among vegetative shoots. The average and individual features stay within the
variability of the species (Kucowa 1971, Á l vare z Fer nánde z 2003). The preliminary
journal 26.indb 258
studies completed to-date fail to support the
existence of a separate taxonomic species, at
least when concerning the features examined
in this project. The differences identified in
the number of inflorescences, diameters of the
capitula and the heights of generative shoots
could also be explained by habitat conditions.
The abundance of nutrients, their quantities,
quality and availability, as well as the quality
of the environment itself are among the fundamental factors affecting the size structure
of individuals in populations. Height is one
of the most variable morphological plant
features observed and modelled in many
species (e.g. Har p e r 1977, 1980, Fa l i ńsk a
1979, Andrzeje wska and Fa lińska 1983,
1986, Mit k a 1988, C z ar ne ck a 1995, 2006,
Łu k as z y ńsk a
1998, Ko st r a k i e w i c z
2008). The differences in biometric traits for
subalpine species were studied, inter alia, in
Veratrum lobelianum Bernh. (Łu k a s z y ńsk a
1998) and Adenostyles alliariae (Gouan) A.
Kern. (By lińska et al. 2000). They resulted
from both climatic (elevation-related) and
edaphic conditions.
D. austriacum seems to prefer open areas, and hence the generative shoots in the
GORCE population are shorter but with higher numbers of leaves and capitula. Strong light
decomposes the elongation hormones, which
results in shorter stems under more intensive
2011-06-28 10:20:22
Phenotypic reaction of Doronicum austriacum
light, whilst at the same time the light enhances assimilation and thus generates larger
quantities of assimilates, which in turn condition a higher number of branches and capitula
(Kop c e w i c z and L e w a k 2005). This adaptation may also be an illustration of the ‘trade
off ’ rule (Fa l ińsk a 1990). Next, the superior
soil properties in the BABIA GÓRA site may
perhaps be suppressed by stronger competition from other macroforb plants (available
ecological niches) and by the shorter vegetation season. These stems are shorter than in
the GORCE population and have fewer numbers of leaves and capitula. The lower number
of leaves in the BABIA GÓRA population is,
however, compensated for by their size. Shadowing and a longer vegetation season resulted
in elongation of stems in the BIAŁA population. The generative individuals growing in
this locality are tall, reaching ca 86 cm on average but produce an average of 2.3 inflorescences and they have lower numbers of leaves.
Shadow and low quantities of nutrients in the
soil may intensify the competition between
individuals in the habitat concerned. When
analysing such a situation in the light of the
‘trade off ’ rule, it can be found than in return
for growing tall, the number of branches is
limited, as the number of inflorescences is.
The reason for such a strategy adopted by the
study Doronicum population could be the periodic adverse environmental conditions. The
expansion of the leopard’s bane in the area occurs at the time of the lowering ground water
table coupled with other profound changes
in habitats (Bróż and Pr zemy s ki 1992,
Wój c i k 1997).
A similar relationship, i.e. the tallest plants
correlated with the lowest number of capitula
on sites with adverse habitat conditions, has
also been observed e.g. in Senecio rivularis
(Waldst. & Kit.) DC. (Czar ne ck a 1995,
2006, 2008). This species grew on dried-up
carr although it prefers moist soil conditions.
And similarly, the density of shoots was very
high. This effect is probably also manifested
in the case of the Doronicum population studied in the BIAŁA site. An additional factor
promoting elongation of shoots is shadowing. More energy spent on growth results in
a smaller number of capitula. Again, a similar
case of the relationship between the height of
shoots and amount of light and soil moisture
journal 26.indb 259
259
has been observed in Iris sibirica L. (Sp orek
and R omb el -Br y z ek 2005, Ko st r a k i e w i c z 2008).
The positive relationship between light
availability and the production of generative
structures has been described in numerous
studies (e.g. D aub enmire 1973, Towp asz
and S z y msk a 1983, And r z e j e ws k a and
Fa l i ńsk a 1986, C z ar ne ck a 1986, 1995,
Fa li ńska 1990). The phenomenon can also
be noted in the Doronicum populations under the study – the highest numbers of capitula were found in the GORCE site, where
the studied population grows in full sunlight.
The capitula there are also the best developed,
namely, in terms of having the largest diameters. The seeds from this site are also the
largest.
Although the Austrian leopard’s bane is
one of the subalpine species, it turned out that
its seeds do not need prior freezing to start
germination. The seeds germinate as well
without such freezing. What is definitely important is the size of seeds, as proven by the
larger seeds from the GORCE site, which germinated faster and where the percentage of
germinating seeds was also higher. Although
the preliminary results indicate a declining
number of germinating seeds with the laps of
time, this hypothesis still needs more precise
verification.
The spatial studies revealed differences
in proportions of vegetative and generative
shoots in particular populations. The number
of vegetative shoots was much higher in the
degraded habitat of the BIAŁA site. It would
be interesting to find why such differences
have occurred there. Probably one of the factors limiting the flower’s development was the
poor quality of the habitat. This hypothesis is
perhaps supported by the fact that after transplanting more than ten young individuals
from the BIAŁA site in early spring (just after the beginning of the vegetation process) to
substitute habitats (home gardens), the flowering plants which appeared in the summer
displayed a greater number of capitula than in
the parental populations (3–5). This number
further increased in subsequent years to 4–7
in 2007, and 4–10 in 2008. Thus, it may be expected that limited flowering, i.e. the increased
proportion of vegetative shoots and the small
number of capitula, resulted from growth un-
2011-06-28 10:20:22
260
Alina Stachurska-Swakoń and Krystyna Kuź
der stress caused by dried-up soil and overshadowing. A number of studies indicate that
there is a certain critical size of an individual,
which determines the beginning of the generative reproduction phase (e.g. Wer ner 1975,
E li áš 1985, Mit k a and Tu m i d aj ow i c z
1993, C zar ne ck a 1995, Aars en and Jord an 2001, O b es o 2002). This critical size
may be conditioned by both ‘calendar’ age and
individual features, or quality of habitats (e.g.
Andrzeje wska and Fa lińska 1983, 1986,
B a l l e g a ard and War ncke 1985, Czarne cka 1986, 2008, Mit k a 1989, Hemb org and Karls on 1998a, b, Fa l ińska 1989,
1990, 1997, Bü h l er and S chm id 2001).
7. CONCLUSIONS
1. Doronicum austriacum demonstrates
a wide ecological amplitude and great tolerance towards adverse (changing) habitat conditions.
2. Habitat and coenotic conditions affect the height, quantitative and biometric features of leaves, and above all, the
number and size of inflorescences. The
proportions of generative and vegetative
shoots and their spatial arrangements
also change.
3. There were statistically significant differences between the populations under
study. The results obtained in the experiment involving transplantation of wild
plants into garden conditions allows for
the assumption that these features are
subject to environmental modification.
4. Despite the fact that D. austriacum is
a mountain species, its seeds do not require freezing.
5. The present level of knowledge does
not support the idea of a morphologically
different lowland form of the species.
ACKNOWLEDGMENT: The authors would
like to thank to the Babia Góra National Park and
the Gorce National Park for the permission to
collect data. Thanks are also due to mgr Barbara
Szczepanowicz for her help with the soil analysis.
The authors are very grateful to Professor Helena
Trzcińska-Tacik and Dr hab. Józef Mitka for critical reading of the manuscript. This work was partially supported by Polish Ministry of Science and
Higher Education, grant No 2P04G 09528.
journal 26.indb 260
8. REFERENCES
Aars en L.A., Jord an C.Y. 2001 – Betweenspecies patterns of covariation in plant size,
seed size and fecundity in monocarpic herbs
– Ecoscience, 8: 471–477.
Aler ic K.M., Kirkman L.K. 2005 – Growth
and photosynthetic responses of the federally
endangered shrub, Lindera melissifolia (Lauraceae), to varied light environments – Am. J.
Bot. 92: 682–689.
A l lendor f F.W., Lui kar t G. 2007 – Conservation and the genetics of populations – Blackwell, 642 pp.
Á lvare z Fer nánd e z I. 2003 – Systematics of
Eurasian and north African Doronicum (Asteraceae: Senecioneae) – Ann. Missouri Bot.
Gard. 90: 319–389.
Andrzeje wska L., Fa lińska K. 1983 – Struktura wielkości osobników w populacji [Structure of individuals size in population] – Wiad.
Ekol. 29: 3–31 (in Polish, English summary).
Andrzeje wska L., Fa lińska K. 1986 – Struktura wielkości osobników w populacji [Structure of individuals size in population] (In:
Populacje roślin i zwierząt. Ekologiczne studium porównawcze [Plant and animal populations. Comparative ecological study], Eds: R.
Andrzejewski, K. Falińska) – PWN, Warszawa, pp. 111–134 (in Polish).
B a l l e g a ard T. K . , War ncke E . 1985 – The age
distribution of a Cirsium palustre population
in a spring area in Jutland, Denmark – Hol.
Ecol. 8: 59–62.
Bróż E., Przemyski A. 1992 – Omieg górski Doronicum austriacum – występowanie,
zagrożenia oraz uwagi dotyczące ochrony jego
reliktowych stanowisk w Polsce [Doronicum
austriacum – its occurence and development,
as well as some remarks on the protection of
its relic localities in Poland] – Chrońmy Przyr.
Ojcz. 48: 51–61 (in Polish).
Bü h l e r C . , S ch m i d B. 2001 – The influence of
management regime and altitude on the population structure of Succisa pratensis: implications for vegetation monitoring – J. Appl. Ecol.
38: 689–698.
By l i ńs k a E . , Gi z a k W. , Matus i e w i c z
O. , Winte r B. 2000 – Ekologia populacji
miłosny górskiej Adenostyles alliariae (Guan)
A. Kern w zróżnicowanych warunkach
edaficznych Karkonoszy [Ecology of Adenostyles alliariae (Guan) A. Kern. population
in various sites conditions in the Karkonosze Mts.] – Opera Corcontica, 37: 175–180
(in Polish).
Czar ne cka B. 1986 – Biological properties of
Majanthemum bifolium (L.) F.W.Schm. poly-
2011-06-28 10:20:22
Phenotypic reaction of Doronicum austriacum
cormones under various ecological conditions
– Acta Soc. Bot. Pol. 55: 659–678.
Czar ne cka B. 1995 – Biologia i ekologia
izolowanych populacji Senecio rivularis (Waldst.
et Kit.) DC. i Senecio umbrosus (Waldst. et Kit.)
[Biology and ecology of the isolated populations of Senecio rivularis (Waldst. et Kit.) DC.
and Senecio umbrosus (Waldst. et Kit.)] – Maria
Curie-Skłodowska University Publisher, Lublin, 263 pp (in Polish).
Czar ne cka B. 2006 – The status of individuals
within a changing plant population and community: the example of Senecio rivularis (Asteraceae) – Pol. Bot. Stud. 22: 155–164.
Czar ne cka B. 2008 – Spatiotemporal patterns
of genets and ramets in a population of clonal
perennial Senecio rivularis: plant features and
habitat effects – Ann. Bot. Fennici, 45: 19–32.
D aub enmire R .F. 1973 – Roślina i środowisko
[Plant and environment] – PWN, Warszawa,
523 pp. (in Polish).
E li áš P. 1985 – Critical rosette size for flowering
in a biennial plant species (Verbascum speciosum
Schrader) – Acta Univ. Agric. (Brno), 33: 435–444.
Fa lińska K . 1979 – Experimental studies of the
reproductive strategy of Caltha palustris L.
populations – Ekol. pol. 27: 527–543.
Fa lińska K. 1989 – Plant population processes
in the course of forest succesion in abandoned
meadows. II. Demography of succession promoters – Acta Soc. Bot. Pol. 58: 467–491.
Fa l ińska K . 1990 – Osobnik – populacja – fitocenoza [Individual – population –phytocoenosis] – PWN, Warszawa, 310 pp (in Polish).
Fa lińska K . 1997 – Ekologia roślin [Plant ecology] – PWN, Warszawa, 453 pp (in Polish).
Har p er J.L. 1977 – Population biology of plants
– Academic Press, London, 857 pp.
Har p er J.L. 1980 – Plant demography and ecological theory – Oikos, 35: 244–254.
Hemb org A.M., Karls on P.S. 1998a – Somatic cost of reproduction in eight subarctic
plant species – Oikos, 82: 149–157.
Hemb org A.M., Karls on P.S. 1998b – Altitudinal variation in size effects on plant reproduction – Ecoscience, 5: 517–525.
Hens en I., Wes che K. 2006 – Relationships
between population size, genetic diversity and
fitness components in rare plant Dictamus albus in Central Germany – Biodivers. Conserv.
15: 2249–2261.
Jas i e w i c z A . 1965 – Rośliny naczyniowe
Bieszczadów Zachodnich [Vascular plants of
the Western Bieszczady Mts.] – Monogr. Bot.
20: 1–336 (in Polish).
Kop c e w i c z J. , L e w a k S . 2005 – Fizjologia
roślin [Plant physiology] – PWN, Warszawa,
807 pp. (in Polish).
journal 26.indb 261
261
Kor naś J., Me dwecka-Kor naś A. 1967
– Zespoły roślinne Gorców. I. Naturalne
i na wpół naturalne zespoły nieleśne [Plant
associations of the Gorce Mts. I. Natural
and semi-natural non-forest associations] –
Fragm. Flor. Geobot. 13: 41–114 (in Polish).
Kos t r a k i e w i c z K . 2008 – Population structure
of a clonal endangered plant species Iris sibirica L. in different habitat conditions – Pol. J.
Ecol. 56: 581–592.
Kos t r a k i e w i c z K . 2009 – The influence of
shadow created by adjacent plants on phenotypic plasticity of endangered species Trollius
europaeus L. (Ranunculaceae) – Pol. J. Ecol. 57:
625–634.
Kovach W.L. 2007. – A Multi Variate Statistical Pakage (MVSP) 3.1. for IBM PC’s. Kovach
Computing Services, Pentraeth, Wales, UK.
Kow a l kows k i A . , Król H . , Ost rows k a A . ,
Sy łek J. 1973 – Instrukcja laboratoryjna
dla pracowni gleboznawczo-nawożeniowych
[Laboratory Guide for Soil Analysis] – Instytut Badawczy Leśnictwa, ZGiN, Warszawa,
228 pp. (in Polish).
Kucowa I. 1971 – Doronicum L. Omieg. (In:
Flora Polska – Rośliny Naczyniowe Polski
i Ziem Ościennych. 12 [Flora of Poland – Vascular plants of Poland and adjacent countries.
12], Eds: B. Pawłowski, A. Jasiewicz) – PWN,
Warszawa – Kraków, pp. 314–319 (in Polish).
L ityński T., Jurkowska H., G orlach E.
1976 – Analiza chemiczno-rolnicza [Chemical
and Agricultural Analyses] – PWN, Warszawa, 330 pp. (in Polish).
Łu k asz y ńsk a I. 1998 – Ekologia populacji Veratrum lobelianum Bernh. w Sudetach [Population ecology of Veratrum lobelianum Bernh.
in the Sudety Mts.] – Acta Univ. Wratisl., Prace
Bot. 77: 125–204.
Matus z k i e w i c z W. 2005 – Przewodnik
do oznaczania zbiorowisk roślinnych Polski
[Guidebook for the Polish plant communities]
– PWN, Warszawa, 537 pp. (in Polish).
May r E. 1974 – Populacje, gatunki i ewolucja
[Population, species and evolution] – Wiedza
Powszechna, Warszawa, 592 pp. (in Polish).
Me dwecka-Kor naś
A.,
Kor naś
J.
Pawłowsk i B., Z arzyck i K. 1977 –
Przegląd ważniejszych zespołów roślinnych
Polski [Review of plant associations in Poland]
(In: Szata roślinna Polski 1 [Plant cover of Poland 1], Eds: W. Szafer, K. Zarzycki) – PWN,
Warszawa, pp. 279–440 (in Polish).
Micha l i k S . , S z ar y A . 1998 – Zbiorowiska
leśne Bieszczadzkiego Parku Narodowego
[Forest communities of the Bieszczadzki National Park] – Monogr. Bieszczadzkie 1: 5–174
(in Polish).
2011-06-28 10:20:22
262
Alina Stachurska-Swakoń and Krystyna Kuź
Miner B.G., Su lt an S.E., Morgan S.G.,
Pa d i l l a D. K . , R e l e y a R . A . 2005 – Ecological consequences of phenotypic plasticity
– Trends Ecol. Evol. 20: 687–692.
Mit ka J. 1988 – Effect of mineral fertilization
upon the forest grass population of Molinia
arundinacea Schrank. I. The use of matrix
model for the simulation of population growth
during ample and insufficient supplies of water – Zesz. Nauk. Uniw. Jagiell. Prac. Bot. 17:
161–174.
Mit ka J. 1989 – Effect of mineral fertilization
upon the forest grass population of Molinia
arundinacea Schrank. II. Population flux and
reproductive effort – Zesz. Nauk. Uniw. Jagiell.
Prac. Bot. 18: 95–112.
Mit ka J. 1997 – Małe izolowane populacje na skraju zasięgu geograficznego - niektóre procesy
ekologiczne i genetyczne [Small, isolated plant
populations at geographical range borders
- some ecological and genetic processes] –
Wiad. Bot. 41: 13–34 (in Polish, English summary).
Mit k a J. , Tu m i d aj ow i c z D. 1993 – Program
ochrony zagrożonych gatunków roślin [Programme for the conservation of endangered
plant species] – Prądnik, 2: 27–37 (in Polish).
O b es o J. R . 2002 – The costs of reproduction in
plants – New Phytol. 155: 321–348.
Oleksynowa K., Tokaj J., Ja kubiec J. 1976
– Metody analityczne analizy gleby [Analytic
methods of soil analysis] (In: Przewodnik
do ćwiczeń gleboznawstwa i geologii dla studentów AR. Część II [Guidebook of soil and
geology laboratory for AR students. Part II],
Ed: T. Komornicki) – Agriculture University
AR Publisher, Kraków, 132 pp. (in Polish).
Pie ch K. 1924 – Doronicum austriacum Jacq.
i Cochlearia officinalis L. w okolicy Olkusza
[Doronicum austriacum Jacq. and Cochlearia
officinalis L. in the vicinity of Olkusz] – Acta
Soc. Bot. Pol. 2: 216–221 (in Polish).
Pig liucci M. 2005 – Evolution of phenotypic
plasticity: where are we going now? – Trends
Ecol. Evol. 20: 481–486.
S ch licht ing C.D. 1986 – The evolution of phenotypic plasticity in plants – Ann. Rev. Ecol.
Syst. 17: 667–693.
S ch licht ing C.D. 2002 – Phenotypic plasticity
in plants – Plant Spec. Biol. 17: 85–88.
Sp orek M., Romb el-Br yzek A. 2005 – Wetland restoration enhances the development of
protected species (Iris sibirica L.) - a case study
– Pol. J. Ecol. 53: 591–595.
St a chursk a - Sw a koń A . 2009 – Plant communities of the Adenostylion alliariae Br.-Bl. 1926
in the Carpathians – Initial results (In: The
role of the geobotany in biodiversity conservation, Eds: J. Holeksa, B. Babczyńska-Sendek,
S. Wika) – Silesian University Publisher, Katowice, pp. 125–134.
Szafer W. 1930 – Element górski we florze
niżu polskiego [Mountain element in the
flora of the Polish lowlands] – Rozpr. Wydz.
Matem.-Przyr. Pol. Akad. Umiejętn., Dz. B.
Nauki Biol. 69, Nr 3: [85]–[196] (separatum:
1–112) (in Polish).
Sz afer W. 1977 – Szata roślinna Polski niżowej
[Vegetation of the Polish lowlands] (In: Szata
roślinna Polski 2 [Plant cover of Poland 2],
Eds: W. Szafer, K. Zarzycki) – PWN, Warszawa, pp. 17–188 (in Polish).
Tow p a s z K . , S z y msk a M . 1983 – Struktura
i dynamika populacji Milium effusum L. w lasach koło Polanki–Haller na Pogórzu Wielickim (Południowa Polska) [Structure and dynamics of populations of Milium effusum L. in
a forest near Polanka–Haller in the Wielickie
Foothills (Southern Poland)] – Zesz. Nauk.
Uniw. Jagiell., Prace Bot. 11: 109–142 (in Polish, English summary).
Va l l a d are s F. , Gi anol i E . , G om e z J. M .
2007 – Ecological limits to plant phenotypic
plasticity – New Phytol. 176: 749–763.
Wer ner P.A. 1975 – Prediction of fate from
rosette size in teasel (Dipsacus fullonum L.) –
Oecologia, 20: 197–201.
Wój ci k W. 1997 – Ekologiczne następstwa
odprowadzania ścieków przemysłowych
zawierających metale ciężkie do ekosystemu
mokradłowego rzeki Białej [Ecological consequence of sewage input containing heavy metals to fen ecosystem of Biała river] – Rozpr.
Monogr. AGH Kraków, 63: 1–111 (in Polish,
English summary).
Z aj ą c A . , Z aj ą c M . 2001 – Distribution atlas
of vascular plants in Poland – Laboratory of
Computer Chorology, Institute of Botany, Jagiellonian University, Cracow, 716 pp.
Received after revision Februrary 2011
journal 26.indb 262
2011-06-28 10:20:23