POLISH JOURNAL OF ECOLOGY

(Pol. J. Ecol.)

61

683691

2013

Regular research paper

Krzysztof OKLEJEWICZ 1, Eugeniusz CHWASTEK 2, Marian SZEWCZYK 3,

Andrzej BOBIEC 4, Jzef MITKA 5*
Department of Botany, University of Rzeszw, Zelwerowicza 4, 35601 Rzeszw, Poland
Bielsko-Biaa Academy of Jzef Tyszkiewicz, Nadbrzena 12, 43300 BielskoBiaa, Poland
e-mail: gienekbiolog@wp.pl
3
Institute of Agriculture, Jan Grodek State Higher Vocational Academy in Sanok, Mickiewicza 21
38500 Sanok, Poland
4
Department of Agroecology, University of Rzeszw, wikliskiej 2A, 35601 Rzeszw, Poland
5
Institute of Botany, Botanical Garden, Jagiellonian University, Kopernika 27, 31501 Krakw, Poland
*e-mail: j.mitka@uj.edu.pl (corresponding author)
1

DISTRIBUTION OF CRATAEGUS (ROSACEAE) IN S-E POLAND

ALONG A GRADIENT OF ANTHROPOGENIC INFLUENCE
ABSTRACT: The impact of land use intensity on the taxonomic composition of hawthorns
in four regions of south-eastern Poland was investigated. As a result of different human settlement histories, two of the regions are 5464%
forested nowadays, and the remaining two regions are 7078% transformed into arable land.
Numerical analysis and Monte Carlo permutation
tests showed that the purely geographical component of the species distribution, linked with
the land use intensity, was statistically significant
(1.97%, P = 0.002), as well the pure site component
(1.17%, P = 0.010). The human-induced opening
of the landscape promotes Crataegus monogyna.
Forests are occupied by C. laevigata, C. rhipidiphylla, C. rhipidophylla Gand. var. lindmanii and
the hybrid C. macrocarpa (C. rhipidophylla C.
laevigata); on forest edges the hybrid C. media
(C. monogyna C. laevigata) tends to occur. Crataegus subsphaericea (C. monogyna C. rhipidophylla) is rare and tends to occur in thickets, and
a triple hybrid C. monogyna C. rhipidophylla
C. laevigata colonizes recently abandoned fields.
Hybridization seems to be an efficient evolutionary strategy of hawthorns in the face of humaninduced transformations of the landscape.
KEY WORDS: CORINE Land Cover, hawthorn, human impact, hybrids, landscape transformation, land use, species distribution

journal 36.indb 683

1. INTRODUCTION
Evolutionary processes in human-dominated landscapes are poorly understood, however, there is little doubt that human activities
have altered them. They are best known from
the population-genetic studies. For example,
Pe rg ams and L a c y (2008), using museum
and recently collected specimens, found rapid
morphological and genetic (mtDNA) change
in a population of white-footed mice in the
Chicago region during the past 150 years.
The other problem, to which little attention
is paid, is the issue of the consequences of
changing evolutionary trajectories of species
that persist in human-altered environments
(Sm it h and B e r nat he z 2008). Human
impact is one of the most important factors
changing the vegetation cover of the Earth,
and results in synanthropization processes
(Kor na 1981, 1983). Some plants increase
their ranges (hemerophiles), and at same time
others decrease as an effect of human impact
(hemerophobes). The most interesting group
of taxa, is one in which the evolutionary response to human alterations is especially
rapid. It forms a group of flowering plants
with a flexible type of asexual reproduction,
i.e. apomixis (C z api k 1996). To this group

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Krzysztof Oklejewicz et al.

belong, among others, Taraxacum dandelions, Hieracium hawkweeds, Rubus brambles

or blackberries, and Crataegus hawthorns. In
hawthorns evolutionary change may be rapid
and flexible due to pseudo-gamous gametophytic apomixis (Ta lent 2009). Also, rapid
ploidy-level changes are possible if a meiotically reduced egg cell develops by parthogenesis, or if a meiotically unreduced egg cell is
fertilized. These phenomena are almost always associated with hybridization (Ta le nt
and Dick ins on 2005, L o et al. 2009).
In Europe, hawthorns were taxonomically revised by C hr istens en (1992, 1997).
In south-eastern Poland three basic taxa have
been found: C. monogyna Jacq., C. rhipidophylla Gand. and C. laevigata (Poir.) DC., and three
related nothospecies: C. subsphaericea Gand.
(= C. monogyna C. rhipidophylla), C. macrocarpa Hegetschw. (= C. rhipidophylla C.
laevigata) and C. media Bechst. (= C. monogyna C. laevigata). Additionally, we included
an extreme morphological form of hawthorn
represented by C. rhipidophylla Gand. var. lindmanii (Hrabtov) K. I. Chr., as well as forms
described as a triple hybrid C. monogyna C.
rhipidophylla C. laevigata (Holub 1992).

Generally, information on the ecological preferences and amplitudes of the species and
nothospecies of hawthorns is absent.
The genus Crataegus is known for its polyploidy and gametophytic apomixis. Apomixis
and reproductive barriers between cytotypes
are factors that reduce the frequency of gene
flow among populations, and may ultimately
led to allopatric speciation (L o et al. 2009). In
this way, agamic complexes form evolutionary potential to promote new genetic variability and ecological varieties. D i ck i ns on
et al. (2008) pointed out that human disturbance of the landscape may have diversified
originally diploid sexual lineages as a result
of polyploidy and agamospermy, including
reticulate evolution (L o et al. 2010).
The aim of this paper is to reveal the general ecological profiles of the hawthorns in
four regions of southern Poland, with different histories of human settlement. Our main
hypothesis is that the taxonomic differentiation of hawthorns in the particular regions is
associated with the different patterns of land
use. The relatively undisturbed region of the
Beskid Niski Mts with the youngest settlement history and large share of forest areas,

Fig. 1. Localization of the four study regions in south-eastern Poland in the ATPOL grid. A map courtesy of A. Zajc and M. Zajc, Laboratory of Computer Chorology, Institute of Botany, Jagiellonian
University.

journal 36.indb 684

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Crataegus in man-made landscape

685

Fig. 2. The land use categories according to the CORINE Land Cover database in the regions under
study. The figure does not consider wet areas and water bodies that form altogether 0.02% in the Beskid
Niski and 1.5% in the Pogrze Przemyskie.

should be dominated by a specific group of

species. On the other hand, the region of the
Przedgrze Rzeszowskie, which has been disturbed by human intervention, has to be supposedly characterized by the occurrence of
the hybridogenous species of the genus. Specifically, we expect that each of the regions
with a unique land use pattern has different
share of hawthorn species. The main premise
of the studies is that the origin and spread of
hybridogenous species was facilitated by the
opening of the landscape.
2. STUDY AREA AND METHODS
The hawhtorns were collected in three
physiographic units within the Polish Western
Carpathians: the Beskid Niski Mts, Pogrze
Przemyskie (Przemyskie Foothills), and
Pogrze Strzyowskie (Strzyowskie Foothills). The specimens were dried, attached to
paper sheets and labelled in a standard manner, and deposed in the herbarium of Institute
of Botany, Jagiellonian University (KRA). The
fourth physiographic unit, the Przedgrze
Rzeszowskie (Rzeszowskie Foothills) are located the most northwards and belongs to the
Kotlina Sandomierska (Fig. 1). Here, mainly
podzolic soils occurs, in contrast to the brown
soils in the Carpathian area.
The Beskid Niski and Pogrze Przemyskie
regions are characterized by large complexes
of forests and relatively well preserved natu-

journal 36.indb 685

ral plant communities. The Beskid Niski is

the most diverse area in terms of topography and geology. It includes the highest altitudes, rich relief and many variants of flysch
bedrock (B a l on et al. 1995). The Pogrze
Strzyowskie is characterized by deep and
narrow folds formed by the soft geological
substratum creating great site diversity. It is
a rural area with extensive farming and scattered forests. The Przedgrze Rzeszowskie
is the most distinct of the three areas and is
characterized by small height differences,
high human impact and almost total deforestation (Fig. 2). In the Pogrze Przemyskie
(especially its northern part) and eastern part
of the Przedgrze Rzeszowskie the linear pottery communities are dated back to the Neolithic times (C z ekaj - Z ast aw ny 2008). In
historical times they were settled relatively
early, in the 14th century. The second wave of
settlement, dated to the 15th17th centuries, is
related to the colonization of the Beskid Niski
(L a ch 1968, G rk a 1995).
The area under study is differentiated according to the historical and present impact
of man. The Beskid Niski belongs to one of
the most forested regions in Central Europe.
Forests cover 64.5% of its area (Fig. 2). In the
19th century overpopulation of the area of the
Beskid Niski led to extensive deforestation.
The process ceased after 1945, triggering successional changes leading to the recovery of
forest communities on abandoned meadows

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Krzysztof Oklejewicz et al.

and fields (L ach 1968, L ach and Z i t ar a

1989). Recently, they have been turned to
fallows dominated by invasive plant species
as Solidago canadensis and Heracleum sosnowskyi.
According to the CORINE Land Cover
database, we specified a percentage of the
particular land use categories: Arable land,
Forests, Wet areas, Water bodies, Meadows
and Anthropogenic sites (Fig. 2).
The occurrence of eight taxa of Crataegus
was checked using a systematic-random sampling in all squares 2 2 km of an ATPOL
system (Fig. 1). In total, 1429 records were
entered. We recognized, at each sample site,
the taxon of Crataegus, the type of plant community, exposure and altitude (m above sea
level). In order to secure the methodological
consistency all field data were collected by the
same team and species determinations were
done by one of us (KO).
In the trial project a DCCA analysis was
used for the calculation of the longest underlying gradient in the data set. The longest
length of gradient was 2.662 SD. It was close
to 3.0, a value which delimits the use of the
linear or unimodal model of species response
(L ep and mi l auer 1999). Thus, we used
both models: a linear with Redundancy Analysis RDA and a unimodal ordination method,
namely the Canonical Correspondence Analysis CCA (ter Braa k 1987).
The explanatory data set includes both
the geographical (regions of occurrence, the
CORINE Land Use Categories) and site (type
of plant community, exposure and altitude)
variables (see Fig. 3). The nominal (qualitative) variables were coded as 0/1 dummy
variables. In order to establish how much
variance is explained by the site vs geographical variables, a variance partitioning procedure, as described by B orc ard et al. (1992),
was carried out. By making two canonical ordinations, each of them constrained by one of
the sets of explanatory variables: namely the
site and geographical, one can obtain a measure of the total importance for the species
data of (1) the effects of the site conditions
and (2) the geographical structure. Additionally, some species and site variables may
share a common spatial structuring that has
been calculated by the subtraction of the partial effects of the site and geographical vari-

journal 36.indb 686

Fig. 3. CCA (a) and RDA (b) of Crataegus in

south-eastern Poland with environmental and
geographical variables as explanatory factors of
species distribution. laevig C. laevigata, lin C.
monogyna var. lindmani, monog C. monogyna,
macro C. macrophylla, media C. media,
subs C. subsphaericea, triple triple hybrid.
BNiski Beskid Niski, PPrzem Pogrze Przemyskie, PRzesz Przedgrze Rzeszowskie; AN
anthropogenic sites, including road and rail embankments, AR agricultural areas, BO boggy
and wet sites, FO forest sites, ME meadows,
WO water bodies; Forest forest sites, Edges
forest ecotones, Fallows former agricultural land
and balks, Anthrop anthropogenic sites, including road and rail embankments, Thicket isolated
dense stands of tall shrubs; E east, SW southwest, N north, NE north-east, S south, SE
south-east, W west, NW north-west, ExpNull
no exposure, alt altitude (m above sea level).

ables. The significance of the first canonical

axis and all canonical axes were checked by
a Monte Carlo permutation test unconstrained permutations in a reduced model in
499 runs. The reduced model method better

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Crataegus in man-made landscape

maintains the Type 1 error in small data sets.

The frequency data of the CORINE Landuse
Categories in particular regions were angular transformed according to the formula
x = SQRT ARCSIN (x).
The permutation tests of significance of
all correspondence canonical analysis CCA
axes calculated the Trace statistics, i.e. the
sum of all canonical eigenvalues, in relation
to the total inertia, i.e. the sum of all eigenvalues. In that way the significance of speciesenvironmental relations in the direct analysis
was explained, in 499 random permutations.
In order to evaluate the explanatory power of the particular quantitative and qualitative variables, the forward selection procedure was applied, using the partial Monte
Carlo permutations in 499 runs. At the beginning of the procedure each explanatory
variable was tested separately to estimate its
independent, marginal effect. Then, the best
explanatory variables were consecutively
added to the model and each time the effect
of the added variable was calculated, i.e. its
conditional, partial effect was estimated.
The calculations were carried out with
the use of the CANOCO for Windows software, and the graphics with the use of the
Canodraw for Windows (ter Bra a k and
mi l auer 2002).
3. RESULTS
The Monte Carlo test of significance
of the first canonical axis of CCA (eigenvalue = 0.201) gave F-ratio 42.666

687

and P = 0.002, and significance of all canonical axes (Trace = 0.322) F-ratio = 3.613 and
P = 0.002. Similarly, in RDA the significance
of the first canonical axis (eigenvalue = 0.062)
gave F-ratio = 95.238 and P = 0.002, and significance of all canonical axes (Trace = 0.073)
gave F-ratio = 5.907 and P = 0.002.
We checked the importance of the geographical, as well as the site factors affecting
the distribution of Crataegus species in southeastern Poland. A variance partitioning shows
that both groups of variables explain, statistically significantly, the species variability. The
purely geographical component of the species distribution is negligibly more important
(1.97%, P = 0.002) than the purely site component (1.17%, P = 0.010, Table 1).
The results of CCA and RDA are given in
Figs 3a and b. For CCA the total inertia equals
6.946 and the Trace 0.322 (Table 1). In both
analyses Axis 1 describes a geographical gradient of species distribution among the Beskid
Niski vs Pogrze Rzeszowskie and Pogrze
Przemyskie. The Pogrze Strzyowskie has
a different position, located at Axis 2, and is
characterized by a large share of fallows. According to the CCA analysis, here the triple
hybrid found optimum conditions, RDA
showed rather C. subsphaericea. The forward selection of the explanatory data shows
that the region of the Beskid Niski and occurrence of meadows (ME) contribute mostly
to the species variability (Table 2). Similarly,
SW exposition, Forests, ExpNull, Fallows and
Edges can be considered as important variables. Crataegus monogyna seems to be asso-

Table 1. Partitioning of variation of a species data explained by Environment and by Region. Environmental characteristics include the type of plant community, exposure and altitude. Trace sum of all
canonical eigenvalues; P corresponding probability value obtained by the Monte Carlo permutation
test, 499 random permutations.
Component

a
b
c
d
e

Source

Site+Region
Site
Region
Covariation between b and c
Unexplained

0.310
0.173
0.229
0.092
6.636

f
g
d+e+f+g

Pure Site*
Pure Region**
Total

0.081
0.137
6.946

*Region set as covariable

**Site set as covariable

journal 36.indb 687

Trace

F-ratio
P
% Variance
Analyses without covariables
3.670
0.002
4.46
2.416
0.002
2.49
12.191
0.002
3.30
1.32
95.54
Analyses adjusted for covariables
7.290
0.010
1.17
9.713
0.002
1.97

100

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Krzysztof Oklejewicz et al.

688

Table 2. Ranking environmental variables in importance in the unimodal (CCA) and linear (RDA)
models by their marginal (left) and conditional (right) effects of the ecological factors (see Fig. 3), as
obtained by the forward selection (Lambda1 = fit = eigenvalue with a variable only); LambdaA = additional fit = change in eigenvalue); P = significance level of the effect, Fratio = as obtained with a
Monte Carlo permutations under the full model with 499 permutations. Only variables with P 0.05
are shown. For variables explanation see Fig. 3.

Variable
CCA
BNiski
ME
ExpNull
alt
SW
Fallows
Forests

Marginal Effects
Lambda1

Conditional Effects
LambdaA

F-ratio

0.16
0.05
0.01
0.14
0.01
0.03
0.03

0.002
0.002
0.004
0.002
0.014
0.014
0.040

33.16
7.88
4.93
4.28
2.69
2.68
2.03

BNiski
ME
alt
SW
Forests
Edges

0.05
0.01
0.05
0.00
0.01
0.00

0.16
0.03
0.02
0.02
0.01
0.02
0.01
RDA
0.05
0.00
0.01
0.00
0.01
0.00

0.002
0.002
0.002
0.006
0.010
0.018

68.95
13.83
8.26
4.48
3.42
2.78

Fig. 4. Distribution of Crataegus species and nothospecies in south-eastern Poland along a human-induced ecological gradient of influence based on numerical analysis of 1429 records on the ATPOL grid.
laevig C. laevigata, rhipid C. rhipidophylla + var. lindmanii, monog C. monogyna, macro C.
macrophylla, media C. media, subsph C. subsphaericea, triple triple hybrid.

ciated with the Przedgrze Rzeszowskie having the longest history of settlement and the
largest proportion of arable land. The Beskid
Niski is largely forested; here three taxa, C.
rhipidophylla, C. macro and C. lindmanii
found optimal circumstances.
The most diversified region is the Beskid
Niski, where forests, thickets and forest edges
prevail. Within the region, C. rhipidiphylla
and C. macrocarpa found optimum occurrence. Crataegus monogyna is mainly confined to the Pogrze Przemyskie and Pogrze
Rzeszowskie, in flat areas. Here anthropo-

journal 36.indb 688

genic communities prevail. The Pogrze

Strzyowskie is characterized by the occurrence of recent fallows and the optimum of
occurrence of the triple hybrid. Crataegus
laevigata, C. media and C. subsphaericea
occupy outlier positions, however at the opposite ends of the Axis 2 (Fig. 3).
To sum up, the distribution of Crataegus
in the Western Carpathians is statistically significantly shaped by the geographical component, connected with the various types of
human impact. A clear gradient was found
among two species: C. monogyna and C. lae-

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Crataegus in man-made landscape

vigata agg. that tend to occupy opposite ends

of the human impact gradient. The intermediate position is filled by the hybrid species
(Fig. 4).
4. DISCUSSION
All the hawthorn species in Poland are
syntaxonomically classified as characteristic
of the Rhamno-Prunetea class (Matus z k i ew icz 2006, Z aj c and Z aj c 2009). Floristic studies carried out in the second half of the
20th century in the Western Carpathians also
point to their occurrence on forest edges and
in thickets (Z emanek 1989, Z e manek and
Winnick i 1999, Ok lej e w icz 1993), fallows (Jasie w icz 1965, B ar ab as z - Kras ny
2011), rail embankments and balks (Towp asz 1987). Similarly, intensive botanical
studies in the Pogrze Przemyskie by Wolanin (unpublished) point to a wide spectrum
of hawthorn occurrence in forests, on forest
edges, in thickets, glades, on xerothermic
grasslands overgrown by shrubs, in balks and
fallows and on rail and road embankments.
Crataegus rhipidophylla and C. monogyna
have the widest ecological niche. They are
distributed throughout the whole region under study, however our analyses point to their
demarcation along the ecological gradient.
While Crataegus rhipidophylla is strongly associated with forest communities and forest
edges, C. monogyna is the most frequent in
open anthropogenic sites.
Recently, Crataegus spp. were studied
in Flanders (Belgium) by D epy p e re et al.
(2006). The authors found that, morphologically, C. monogyna and C. laevigata are easily recognized species, however their hybrid
C. media showed a remarkably large range
and overlapped with both parental species
for most of the studied traits. According to
Belgian botanists (cited in D epy p e re et al.
2006) in Flanders, C. media displays better adaptability to changing environments
than C. laevigata. The latter species is rare in
the area under study, but its hybrid is common and occupies an intermediate ecological
niche between the parental species, on forest
edges. The other result of the morphological
investigations concerned C. subsphaerica, a
hybrid between C. monogyna and C. rhipidophylla. In Flanders, its morphology is hardly

journal 36.indb 689

689

distinguishable from C. monogyna, which is

a planted species, found in old hedges to delineate field boundaries and to provide a barrier to grazing livestock. On the other hand,
C. laevigata is less common in Flanders and
it is characteristic for old hedges and wood
banks on richer soil types (D epy p e re et al.
2006). Our results show that C. subsphaericea occurs, similarly to C. monogyna, in regions under strong anthropogenic impact,
where it forms dense thickets. Similarly, another hybrid C. macrocarpa (C. laevigata
C. rhipidophylla) is ecologically close to one
of its parents, i.e. to C. rhipidophylla. In fact,
the parental taxa and its hybrid tend to occur
in forest plant communities.
Hybridization in some plant genera is
quite common. It enhances the degree of
variability in fitness among different hybrid
genotypes. In consequence, hybridization
need not be an evolutionary dead end (R i es eb e rg and C ar ne y 1998). The significance
of hybridization in the context of biodiversity maintenance was critically discussed by
S e ehaus e n et al. (2008). They note that, on
the one hand, hybridization may contribute
to an increase in species numbers by enhancing evolutionary potential, but, on the other
hand, it may effectively reversed speciation.
In fact, our results show that C. laevigata is a
rare species, and its hybrid C. macrocarpa
is much more frequent in the forested Beskid
Niski. It seems that C. laevigata may collapse
in a hybrid swarm here, as a result of reduced
population density, a hypothesis that should
be verified by systematic observational and
experimental studies. Our results strongly support the view expressed by Kor na
(1981) that the breaking of the ecological
barriers brought about by human intervention may lead to creation of new nothotaxa.
The newly originated as an effect of humaninduced open landscapes biotypes (antrophyta anthropogena) are often expansive and
could have given rise to, for example, many
central European meadow species (L and olt
1970, S cholz 1975).
Two hybrid nothospecies: C. subsphaericea and the triple hybrid deserve more attention. They are relatively rare, especially the
triple hybrids, and gravitate towards regions
transformed by humans. The rarity of Crataegus subsphaericea may be explained by the fact

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690

that it is the hybrid between parents with different, non-overlapped ecological profiles. The
question is whether its occurrence in patches of
bushes in the open landscape is merely a random effect, or it is a divergent adaptation of the
hybrid species. Similarly, the occurrence of the
triple hybrid in relatively young fallows in the
Pogrze Strzyowskie also need further observations on the dynamics of the process.
In conclusion, the prolonged, i.e. dating
to Neolithic times, activity of man in southeastern Poland seems to be an important
evolutionary factor shaping the taxonomical
diversity of hawthorns. The opening of the
landscape and the creation of novel habitats
disrupted dispersal barriers and promoted
gene flow between otherwise isolated genotypes and made the hybridization of hawthorns much easier. In effect, new genetic
combinations arose that underwent natural
selection. The process has been continuing
and may have led to the generation of new
hawthorn species, to fit the rapidly changing
anthropogenic sites.
ACKNOWLEDGMENTS: The study was
financially supported by the Ministry of Science
and Higher Education/National Science Centre of
the Poland (N N305 221537). The authors thank
Jerzy Zieliski (Faculty of Forest Botany, University of Life Science, Pozna) for help in determination of Crataegus and Stefan Gawroski (Institute
of Botany, Jagiellonian University) for comments
and literature sources on the history of human
settlement in the Carpathian Foothills.
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journal 36.indb 691

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journal 36.indb 692

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