POLISH JOURNAL OF ECOLOGY
(Pol. J. Ecol.)
59
1
75–85
2011
Regular research paper
Anna ORCZEWSKA*, Małgorzata FERNES
University of Silesia, Faculty of Biology and Environmental Protection, Department of Ecology,
Bankowa 9, 40-007 Katowice, Poland
*e-mail: anna.orczewska@us.edu.pl (corresponding author)
MIGRATION OF HERB LAYER SPECIES INTO THE POOREST
POST-AGRICULTURAL PINE WOODS ADJACENT TO ANCIENT
PINE FORESTS
ABSTRACT: The recovery of species composition typical for ancient forests in recent woods
is a very slow process and may last for decades
or even centuries. It is enhanced only when postagricultural woods are adjacent to ancient ones.
However, even in such a situation of the spatial
contact of both forest types, colonization of recent woods by true forest species is a gradual
process. According to studies focusing on the behaviour of individual species and their colonization rates into recent woods, it can be concluded
that in more fertile habitats the migration process
proceeds faster than on poorer sites. Thus, studies were conducted on light, acidic soils both in
ancient and in adjoining post-agricultural pine
woods (the Dicrano-Pinion Libb. 1933 alliance)
and were focused on the process of the colonization of the herbaceous layer by woodland flora in
recent woods. In eight transects 80 m in length
perpendicular to the ancient/recent ecotone and
consisting of 10 sample plots of 16 m2 laid out
at intervals of 4 m, the percentage cover of herb
layer species was recorded. The migration rates
(based on the occurrence of the farthest individual and on the occurrence of the maximum cover
of a species) for 12 forest species were calculated.
The mean migration rate for all species reached
0.54 m yr–1 when based on maximum cover and
0.67 m yr–1 when based on the farthest individual
and appeared to be lower than those reported in
investigations in more fertile and moister habitats.
The migration rates for individual species ranged
from 0 to 1.21 m yr–1 and were also lower than in
more fertile, black alder woodlands. The migration pattern of Vaccinium myrtillus L., the most
abundant species in pine woods, fits the model
based on the establishment of isolated individuals. The cover of most woodland species increased
with the increasing age of a recent wood. Herb
layer recovery on such sites is slower than in the
more productive, fertile habitats of broadleaved
forests. The ancient and recent pine woods investigated here differed in herb layer species composition despite the secondary succession having
lasted for over 50–60 years.
KEY WORDS: colonization, seed dispersal, recruitment limitation, herb layer recovery,
secondary succession, southern Poland
1. INTRODUCTION
The first records of afforestation on postagricultural sites in Poland date back 180
years (S ob c z a k 1996). However, the period
after the World War II was the first well-documented one in Polish forest management
where a noticeable increase in forest cover
was recorded. This was the result of the afforestation of almost one million hectares of
abandoned land, mainly formerly used for
agriculture (Tus zy ńsk i 1990). Scots pine
(Pinus sylvestris L.) was, and still remains, the
76
Anna Orczewska, Małgorzata Fernes
main species used for afforestation since sites
with poor soils are the first ones where farming practices are abandoned. Subsequently,
such sites are converted into forests.
Forests of a post-agricultural origin initially do not contain woodland species in
their herb layer. The natural recovery of species composition typical for ancient forests
in recent woods isolated from the ancient,
source woodlands is a very slow process and
may last for many centuries (Peterken 1977,
Peterken and Game 1984, Fa lińsk i 1986,
Mat l a ck 1994). This is due to the fact that
most woodland herbs have a very poor dispersal potential (Dzwon ko and L oster
1992, B ossuyt et al. 1999, B ossuy t and
He r my 2000). Colonization of recent woods
by woodland flora is enhanced only when
post-agricultural woods are adjacent to ancient ones (Peterken and Game 1984, Dz won ko and L oster 1992, Dzwon ko 1993,
Dzwon ko and Gaw rońsk i 1994, Mat l a ck
1994, Br unet and von Oheimb 1998a, b,
B ossuyt et al. 1999, B ossuyt and He r my
2000, Dzwon ko 2001a, b). However, even in
such a situation of the spatial contact of both
forest types, colonization of recent woods by
true forest species is a gradual process. The
poor dispersal abilities of woodland plants
and their lack of the ability to compete with
vigorously growing species with a wide ecological amplitude are responsible for the slow
rate of herb layer recovery in recent woods
(Her my et al. 1999, Honnay et al. 2009).
Although species traits are among the main
reasons for the slow rate of herb layer development in recent woods, other factors are also
responsible for the pace of that process. For
example, herb layer restoration is influenced
by the dominant tree species in post-agricultural woods. Recent woods can be effectively
colonized by woodland flora when suitable
habitats for the germination and establishment of forest plants exist. Trees for which
the leaf litter quickly decomposes provide
much better conditions for herb layer restoration than stands composed of species with
slowly decomposing litter (Br unet and von
Oheimb 1998a, Dzwon ko 2001a, b, O rcze w ska 2009). According to studies focusing on the behaviour of individual species and
their colonization rates into recent woods, it
can be concluded that in more fertile habitats
the migration process proceeds faster than
on poorer sites (Br u ne t and von Ohe i mb
1998a, Dz won ko 2001b, O rc z e w sk a
2009). Papers describing the pace of colonization of recent forests by ancient woodland
flora in Europe refer to broadleaved, usually
oak-hornbeam communities, recent stands
with black alder, or to pine plantations located on the habitats of deciduous forests (Dzwon ko and L o ste r 1992, Dz won ko 1993,
Dz won ko and G aw rońsk i 1994, Br u ne t
and von Ohe i mb 1998a, b, B o ssuy t and
He r my 2000, Dz won ko 2001a, b, Wu l f
and He i n ke n 2008, O rc z e w sk a 2009,
2010). Although Pinus sylvestris is the most
common species used in afforestation, recent pine stands have not been the subject
of colonization studies except for the investigations by Dzwon ko (2001a, b) and Wu lf
and Hei n ken (2008), which focused on pine
stands on the habitats of rich, broadleaved
forests. Thus, the reason for undertaking this
research was the absence of information concerning the process of herb layer recovery in
recent pine woodlands growing on the poorest agricultural sites.
The main aim of the survey was to calculate the rates of migration of forest species
into post-agricultural pine woods located in
direct proximity to ancient pine forests (Dicrano-Pinion alliance) and to check whether
the migration potential of species is related
to their dispersal modes. The investigation
also helped to answer the question of whether the process of the colonization of recent
pine stands by woodland flora proceeds more
slowly on such poor sites compared to recent
woods in richer and wetter habitats.
2. STUDY AREA
The study was conducted in four recent
pine woods adjacent to ancient pine forests in
the Opolska Plain and Woźnicki Escarpment,
two neighbouring regions of southern Poland
(Table 1). The status of ancient forests was
confirmed since they are present in the old
cartographic sources by Wieland-Schubart,
dated 1739, and on successive maps from
1880. Thus, they have been part of the landscape for over 260 years. Recent pine woods
were planted after the World War II on sites
which had formerly been managed as arable
Migration of herb species into recent pine woods
77
Table 1. Detailed information about the study sites.
No. transect Geographical position
o
o
Age of
recent Geographical region
wood
I, II
50 34’N; 18 54’E
25
III, IV
50o50’N; 18o16’E
34
V, VI
50o47’N; 18o20’E
55
VII, VIII
50o33N; 18o55’E
65
Forest type
Leucobryo-Pinetum W. Mat. (1962) 1973/
Opolska Plain
Querco roboris-Pinetum (W. Mat. 1981) J.
[Równina Opolska]
Mat. 1988
Opolska Plain
[Równina Opolska]
Leucobryo-Pinetum W. Mat. (1962) 1973
Leucobryo-Pinetum W. Mat. (1962) 1973/
Opolska Plain
Querco roboris-Pinetum (W. Mat. 1981) J.
[Równina Opolska]
Mat. 1988
Woźnicki Escarpment
[Próg Woźnicki]
Leucobryo-Pinetum W. Mat. (1962) 1973
fields, but subsequently replaced by pastures
or meadows. Due to this fact there is a high
probability that no seeds of woodland herbs
remained in the soil seed bank of the postagricultural pine woods. Both forest types
grow on a flat area on albic arenosols, haplic
arenosols, and gleyic podzols (nomenclature
after WRB 1998). The forests are owned and
managed by the Polish State Forests.
3. MATERIAL AND METHODS
3.1. Data collection
Fig. 1. Transects used for data collection. Quadrats (4  4 m) were surveyed along eight transects
across ancient-recent forest ecotones. Ancient
forest plots were marked with ‘–’, whereas ‘+’ indicated recent woodland plots. Ecotone plots were
marked as Ea.
In June–July of 2003 two transects per
stand approximately 80 m in length and laid
out perpendicularly across the boundary
between the ancient and recent pine woods
were set up. Ancient forests representing either the Leucobryo-Pinetum W. Mat. (1962)
1973 or a community with features intermediate between Leucobryo-Pinetum and Querco
roboris-Pinetum (W. Mat. 1981) J. Mat. 1988
bordered recent pine stands of the ages of 25,
34, 55 and 65 years. Each point of the transects in their recent woodland part was situated closer to the boundary with the ancient
woodland being studied than to any other
forest of such an origin. Each of the transects
consisted of 10–11 plots, 4  4 m in size laid
out at intervals of 4 m. Four plots (28 m in
overall transect length) were situated in the
ancient forest and six (in total 44 m) in the recent wood. When the ecotone zone was equal
or wider than 4 m, an additional plot adjoining ancient forest edge was set up (Fig. 1). In
78
Anna Orczewska, Małgorzata Fernes
total 32 quadrats in the ancient forests, 48 in
the recent wood and three in the ecotone were
studied. In each plot the percentage cover of
vascular plant species (1, 5, 10%, and then at
10% intervals) was estimated. Additionally,
the spatial distributions of Vaccinium myrtillus and V. vitis-idaea, the two most widely occurring ancient woodland species typical for
pine woods, were mapped and presented in
a graphic form. In order to show their distribution and horizontal structure in the whole
length of the transect, they were mapped both
in the sampling and in interval plots.
3.2. Data analyses
The Fisher exact probability test was
used to compare species frequency in the
two woodland types in order to detect the
herbs recorded significantly more often in
ancient pine woods. Then, the migration
rates (m yr–1) for all of the species predominantly occurring in ancient woodland sites
were calculated. Species listed as ancient
woodland indicators for Poland, according
to Dz won ko and L o ster (2001), were also
included in the set of herbs for which the migration rates were calculated. This resulted in
12 species for which these calculations were
done. The procedure followed the one proposed by Mat l ack (1994). Thus, the migration rates were based on the occurrence of the
farthest individuals, and when possible, also
on the most distant occurrence of the species’
maximum cover. In order to compare all of
the recent woods, regardless of the difference
in age of their stands, standardized distances
were taken into account:
MR = d × a–1
(1)
where MR is the migration rate, m yr–1, d
is the occurrence of the farthest individual/
maximum cover (in meters) and a is the age
of the recent forest (in years). The migration
rates were calculated separately for each transect where the species was recorded and then
the mean rates for a species were counted.
These calculations were limited to those species which were present in at least five plots.
The cover of species in ancient and recent woods was compared using the MannWhitney U-test (Statistica 8.0 package). In
addition, the mean percentage cover of each
woodland species in recent woods of different ages was compared.
Detrended correspondence analysis
(DCA) (Canoco 4.5 program) in its standard
form was used to identify the main directions
of species variation in ancient and recent
woods (species which occurred on a minimum of three plots were taken into consideration).
4. RESULTS
Among the total number of 84 species
recorded in both forest types, V. myrtillus, V.
vitis-idaea, Festuca ovina, Maianthemum bifolium, Pteridium aquilinum, and Deschampsia flexuosa were associated with ancient pine
woods (according to the results of the Fisher’s
exact probability test). M. bifolium and P. aquilinum were present exclusively in ancient
woodland sites, whereas four other species
colonized recent woods, although they differed in their migration capacity (Table 2).
Moreover, within the group of 13 herbs regarded as ancient woodland indicators for
Poland (Dz won ko and L o ste r 2001), Viola
reichenbachiana (like the already mentioned
M. bifolium and P. aquilinum) did not migrate into recent pine woods, whereas other
species showed a diverse colonization potential. In most cases the migration rate for the
farthest individual was much lower than 1 m
yr–1 indicating that, in general, woodland species present in recent woods have a low colonizing capacity. V. myrtillus, Equisetum sylvaticum, and Dryopteris carthusiana are the
exceptions as their migration rates exceeded
1 m yr–1. In addition, Oxalis acetosella was
effective in the colonization of recent stands
since its migration rate reached almost 1 m
yr–1 (Table 2). From the group of 12 species
present in recent pine woods for which the
migration rates were calculated O. acetosella,
Moehringia trinervia, D. carthusiana, and V.
myrtillus occurred in many sites in the whole
area of the recent woods, regardless of the
distance of the plots from the ancient wood
(10 plots of 4 transects; 38 plots of 8 transects, 27 plots of 6 transects, and 26 plots of
7 transects, respectively). They colonized recent woods relatively quickly. Thus, the migration rates for those species underestimate
Auto
Baro
Endo
Endo
Oxalis acetosella L.
Trientalis europaea L.
Vaccinium myrtillus* L.
Vaccinium vitis-idaea* L.
Non migrating ancient woodland species
Maianthemum bifolium* (L.) F. W.
Endo
Schmidt
Pteridium aquilinum* (L.) Kuhn
An1
Viola reichenbachiana Jord. ex
Myrm
Boreau
An2
Myrm
Mycelis muralis (L.) Dumort.
Moehringia trinervia (L.) Clairv.
Epi
Festuca ovina* L. s. str.
0
0
0
10
2
3
27
25
6
19
30
20
26
5
16
10
1
3
6
8
38
14
5
2
17
1
An1
Myrm
7
An1
Luzula pilosa (L.) Willd.
19
Epi
Deschampsia flexuosa (L.) Trin.
Dryopteris carthusiana (Vill.) H.
P. Fuchs
Equisetum sylvaticum L.
Ancient Recent
plots
plots
n=32
n=48
1
6
Frequency
An1
Dispersal
mode
Athyrium filix-femina (L.) Roth
Species
0.0
0.0
0.0
0.16
0.91
0.66
0.87
/
0.59
/
0.36
1.45
0.92
0.53
/
MC
0.0
0.0
0.0
0.61
>1.21
0.79
>0.96
0.82
>0.78
0.4
0.36
1.45
>1.1
0.75
0.78
FI
Pine woods
0.44
–
0.33
–
–
–
0.26
–
–
–
–
–
0.38
–
0.25
MC
0.67
–
0.34
–
–
–
0.43
–
–
–
–
–
0.44
–
0.42
FI
B & O 1998b
0.24
–
–
–
–
–
–
>0.53
>0.53
0.38
–
–
>0.53
–
–
FI
D 2000b
–
–
–
–
–
–
0.65
–
0.82
–
–
–
–
–
–
MC
0.01
–
1.33
–
–
–
0.81
–
2.24
–
–
0.14
1.48
–
1.31
FI
O 2009 o-h
0.00
–
–
–
–
–
0.75
–
0.76
–
–
0.00
0.30
–
–
MC
0.30
–
–
–
–
–
0.66
–
1.07
–
–
0.00
1.59
–
–
FI
O 2009 a-a
–
–
–
–
–
–
0.65
–
1.51
–
–
–
0.88
–
0.17
MC
–
–
–
–
–
–
1.09
–
1.95
–
–
–
1.63
–
1.03
FI
O 2009 w-a
Table 2. Frequencies of woodland species (* – significantly different at P <0.05 based on the Fisher exact probability test) and their mean migration rates (m yr–1) based on the distance of maximum cover of a species (MC) and on the location of the farthest individual (FI). For comparisons, migration rates calculated by other authors are given: B & O 1998b
– Br unet and von Oheimb 1998b, D 2001a – Dzwon ko 2001a, D 2001b – Dzwon ko 2001b, O 2009 o-h – Orcze wska 2009, oak-hornbeam forest habitat, O 2009 a-a – Orcze w ska 2009, alder-ash carr, O 2009 w-a – O rcz e wska 2009, wet alder wood. Dispersal modes are given after Van de r Pijl (1982): Myrm – myrmecochores; Auto – autochores;
Baro – barochores; Epi – epizoochores; Endo – endozoochores; An1 – unwinged anemochores with small diaspores; An2 – anemochores with diaspores with pappus or wings. The ‘/’
sign indicates that in most cases a species occurred with a very low percentage cover, thus not allowing for calculations of the migration rate based on the species’ maximum cover,
whereas the ‘–’ mark means that a species was absent in a particular study. Bold fonts signify ancient woodland indicator species (after Dzwon ko and L o ste r 2001).
Migration of herb species into recent pine woods
79
80
Anna Orczewska, Małgorzata Fernes
Table 3. Mean cover of woodland species in ancient and recent pine woods and the significance level
of the differences between these values according to the Mann-Whitney U-test. n.s. – not significant.
* – species with a higher frequency in ancient pine woods according to the Fisher exact probability test.
Species
Athyrium filix-femina
Deschampsia flexuosa*
Dryopteris carthusiana
Dryopteris filix-mas (L.) Schott
Equisetum sylvaticum
Festuca ovina*
Geum urbanum L.
Luzula pilosa
Maianthemum bifolium*
Melampyrum pretense L.
Moehringia trinervia
Mycelis muralis
Oxalis acetosella
Pteridium aquilinum*
Trientalis europaea
Vaccinium myrtillus*
Vaccinium vitis-idaea*
Viola reichenbachiana
Mean cover of species (%) in
ancient wood
recent wood
0.2
0.8
11.6
4.1
0.7
4.3
0.0
0.1
0.0
0.4
5.3
1.0
0.2
0.02
0.3
0.06
1.8
0.0
0.2
0.04
1.4
0.8
0.2
0.1
6.8
8.3
11.1
0.0
4.9
3.3
29.7
6.6
3.8
0.5
0.1
0.0
P level
n.s.
0.019
0.000
n.s.
n.s.
0.006
n.s.
n.s.
0.002
n.s.
n.s.
n.s.
n.s.
0.018
n.s.
0.000
0.000
n.s.
Table 4. Mean percentage cover of selected woodland plants in recent woods of different ages.
Species
Age of recent wood (years)
Athyrium filix-femina
–
Deschampsia flexuosa
1.8
34
1.3
7.3
Dryopteris carthusiana
4.9
2.9
3.8
5.5
Festuca ovina
1.7
Luzula pilosa
0.1
Oxalis acetosella
0.1
0.1
0.9
0.4
–
–
0.7
1.9
3.3
1.8
–
–
16.7
3.3
–
–
2.5
0.2
15.9
8.0
22.2
–
Trientalis europaea
Vaccinium myrtillus
Vaccinium vitis-idaea
25
their colonization potential. All of the species
which were more frequent in ancient pine
woods also reached higher mean cover in
such woods (Table 3). By contrast, the cover
of D. carthusiana in recent woods was distinctively higher than in ancient woodland sites.
A similar performance was recorded in Athyrium filix-femina, D. filix-mas, E. sylvaticum,
and O. acetosella, but those differences were
not significant (Table 3). The cover of some
woodland plants differed and changed with
time. Many of them were more abundant in
stands of the older age classes compared to
the younger woods (Table 4).
55
2.1
–
65
–
7.3
The spatial distribution of V. myrtillus and
V. vitis-idaea illustrates the general trend in
their abundance in ancient and recent woodland sites (Fig. 2). As expected, both species reached a much higher cover in ancient
woods, although in general V. vitis-idaea was
less abundant than the other species. Only in
two cases (transects V, VI, Fig. 2), where Rubus fruticosus reached a high cover, was the
cover of V. myrtillus in ancient woodland distinctively lower compared to the other sites.
The horizontal structure of the populations of
V. myrtillus and V. vitis-idaea in recent woods
changed with the age of the recent wood, and
Migration of herb species into recent pine woods
I
II
III IV
V
VI VII VIII
81
– are concentrated together with the recent,
non-woodland species. Thus, although some
species from the groups overlap with those
belonging to other groups, the first axis may
represent a gradient from an ancient to recent
wood.
5. DISCUSSION AND CONCLUSIONS
Fig. 2. Spatial distribution of Vaccinium myrtillus
and V. vitis-idaea along the ancient/recent woodland gradient. I–VIII – numbers of transects; for
more information about the transects see Table 1.
their cover was higher in older stands compared to the younger woods (Fig. 2, transects
I–IV vs. VII–VIII).
The amount of variation in species composition in ancient and recent woods explained by the first and second DCA axes
reached 22% (first axis = 11.4). In the scatter
plot of species (Fig. 3) woodland species (left
side, black triangles) are reasonably well separated from the species characteristic for recent woods (right side, grey circles). Furthermore, woodland species which were more
frequent in recent woods (right side, black
squares) – A. filix-femina, D. carthusiana, M.
trinervia, E. sylvaticum and Mycelis muralis
Despite the fact that the ancient pine
woods investigated develop on light, acid
soils and are characterized by fewer species
than mesophilous, deciduous forests, they
still have a set of species occurring predominantly or even exclusively within their herb
layer. Furthermore, even in the situation of
the direct proximity of ancient and recent
pine woods, differences in the composition
of their herb layer species are present. Typical woodland species differed in their performance along the ancient-recent forest gradient. Three herbs did not migrate into recent
pine woods (M. bifolium, P. aquilinum, and
V. reichenbachiana), whereas other species
colonized them very slowly (F. ovina, Luzula
pilosa, and V. vitis-idaea). On the other hand,
V. myrtillus migrated at a pace exceeding 1 m
yr–1. Similar values were obtained for D. carthusiana, E. sylvaticum, and for O. acetosella
(Table 2). The last two species mentioned,
although classified as ancient woodland indicators (Dz won ko and L oste r 2001),
occurred more often in recent pine woods
(Table 2). Furthermore, one of them – D. carthusiana – reached significantly higher cover
in recent woods (Table 3). It appears that for
D. carthusiana neither dispersal nor recruitment limitation play a role, and therefore, it
can be regarded as a good colonizer of recent pine woods. O. acetosella and E. sylvaticum are the two species that mainly occur in
relatively fertile forest habitats, and are present in many European lists of ancient woodland indicators (He r my et al. 1999). However, in the case of the pine forests studied,
they found much better conditions in recent
stands where a combination of previously fertilized soils and the lack of competition from
woodland species like V. myrtillus are probably responsible for the successful establishment of their populations.
Although the set of species for which it
was possible to calculate the migration rates
82
Anna Orczewska, Małgorzata Fernes
Fig. 3. Species ordination plot in ancient and recent pine woods along the first two DCA axes (λ1 = 0.69,
λ2 = 0.65, gradient length = 4.28) on the basis of percentage cover of all species occurring in at least
three plots. – woodland species mainly associated with ancient woods; – woodland species which
were more frequent in recent woods;
– species characteristic for recent woods or present in both
forest types. Species abbreviations are based on three letters of the genus and four of the species: des
flex – Deschampsia flexuosa, fes ovin – Festuca ovina, luz pilo – Luzula pilosa, mai bifo – Maianthemum
bifolium, oxa acet – Oxalis acetosella, pte aqui – Pteridium aquilinum, tri euro – Trientalis europaea, vac
myrt – Vaccinium myrtillus, vac viti – Vaccinium vitis-idaea, ath fili – Athyrium filix-femina, dry cart
– Dryopteris carthusiana, equ silv – Equisetum sylvaticum, moe trin – Moehringia trinervia, myc mura
– Mycelis muralis, agr cani – Agrostis canina, agr capi – Agrostis capillaris, bet pend – Betula pendula, bet
pube – Betula pubescens, cal arun – Calamagrostis arundinacea, cal vill – Calamagrostis villosa, car briz –
Carex brizoides, car oval – Carex ovalis, car pall – Carex pallescens, car pseu – Carex pseudocyperus, des
caes – Deschampsia caespitosa, fag sylv – Fagus sylvatica, fra alnu – Frangula alnus, gal pube – Galeopsis
pubescens, gal vern –Galium vernum, hol moll – Holcus mollis, jun cong – Juncus conglomeratus, jun effu
– Juncus effusus, lys vulg – Lysimachia vulgaris, mel prat- Melampyrum pratense, mol coer – Molinia coerulea, oxa font – Oxalis fontana, pic abie – Picea abies, pin sylv – Pinus sylvestris, poa triv – Poa trivialis,
pop trem – Populus tremula, pot erec – Potentilla erecta, que robu – Quercus robur, que rubr – Quercus
rubra, rub frut – Rubus fruticosus, sci sylv – Scirpus sylvaticus, Sil vulg – Silene vulgaris, sol giga – Solidago
gigantea, sor aucu – Sorbus aucuparia, ste gram – Stellaria graminea, ste medi – Stellaria media, urt dioi
– Urtica dioica, vic crac – Vicia cracca, vio palu – Viola palustris.
is not large (ancient pine woods on sandy,
acid soils have an herbaceous layer of a naturally low species richness), it appears that
dispersal limitation is not the only factor
hampering the colonization of recent woods
by forest species. Some species with effective dispersal modes (P. aquilinum, F. ovina)
either did not migrate to recent pine woods
or were very poor colonizers, whereas other
plants, with poor dispersal abilities, migrated
and established their populations in recent
woods successfully (M. trinervia). Such results are in accordance with the observations made by Si ng le ton et al. (2001) who
Migration of herb species into recent pine woods
reported that the species colonization capacity is not necessarily related to the dispersal
mode of a species. The group of poor colonizers also includes plants with adaptations for
long-distance migration, whereas many herbs
effectively colonizing recent woods are myrmecochorous. P. aquilinum and F. ovina seem
to be recruitment limited. On the other hand,
such species as V. reichenbachiana or L. pilosa,
both being ant-dispersed, are dispersal limited. The lack of M. bifolium in recent woods is
in accordance with the observations reported
by other authors, e.g. Honnay et al. (1998),
who classified it as strongly dispersal limited,
an extremely bad colonizer. One may expect
that its performance may be similar to that of
Convallaria majalis, described by B ossuy t
et al. (1999) in Belgium. Although C. majalis
has ingested berries, according to these authors, as an herbaceous plant of a small size, it
has a reduced migration capacity. Therefore,
its colonization pattern is more like that of a
barochorous species. According to the studies
by Honnay and B ossuyt (2005) and Hon nay et al. (2006) M. bifolium exhibits a limited sexual reproduction capacity and prolonged clonal growth. Thus, its dispersal via
seeds is questionable or at least does not play
an important role in its colonization potential. Furthermore, its ability to spread vegetatively through its rhizomes (a phalanx growth
form) did not help it to colonize recent pine
woods. Thus, the richness and diversity of
woodland species in the herb layer of recent
forests is the combined result of dispersal and
recruitment limitation. In addition, as has
been emphasized by many authors, for example by Her my et al. (1999), Dz won ko
and L oster (2001), Her my and Verhe yen
2007, Honnay et al. (2009), and O rcz ewska (2010), other simple species traits, like
life form, life strategy or habitat requirements
are among the driving factors influencing the
diverse behaviour of a species. Two examples are M. bifolium and V. reichenbachiana,
stress-tolerant species that do not migrate
into recent pine woods. M. bifolium has been
reported as an extremely poor colonizer in
many surveys. Its short characteristics have
already been given here. Both species belong
to the group of small perennial herbs with
heavy seeds (according to the classification
of the emergent groups after Ve rhe ye n et
83
al. 2003) which are generally unsuccessful in
colonizing post-agricultural woods throughout Europe, even in relatively dense forested
landscapes (D e Frenne et al., submitted).
In post-agricultural forests the cover of
many woodland species recorded in ancient
pine woods has grown with the increasing age
of recent woods (Table 4, Fig. 2). The spatial
distribution of V. myrtillus, the most abundant species from that group, seems to fit the
colonization model described by Mat l ack
(1994). According to this model a species colonizes a recent wood by the establishment of
isolated individuals with a subsequent population increase filling in the gaps between
them.
The mean migration rate estimated for
the pine woods investigated reached 0.54 m
yr–1 (the result based on maximum cover)
and 0.67 m yr–1 when based on the farthest
individual, whereas in pine plantations growing on a habitat of deciduous forests in southern Poland the rate reached 0.18 m yr–1–0.38
m yr–1, respectively for mean migration rates
based on maximum cover and on the farthest
individual (Dzwon ko 2001b). In deciduous woodlands in southern Sweden the herb
layer restoration proceeds at 0.3–0.5 m yr–1
(Br u ne t and von Ohe i mb 1998b). On
similar habitats of broadleaved, mesotrophic
forests in central Belgium it was calculated
at 0.5–1.0 m yr–1 (B ossuyt et al. 1999). The
highest rates were reported in the case of recent black alder woods, which are the wettest and most fertile habitat types. Thus, in
typical wet alder woods the mean migration
rates reached 1.20–1.60 m yr–1, for the habitat
of oak-hornbeam communities 1.17–1.63 m
yr–1, and in alder-ash carrs 0.79–1.26 m yr–1
(O rc z e w sk a 2009).
It the case of pine woods it is difficult to
use the term ‘effective colonizer’ when only
rough estimations of the migration rate itself are taken into account. This is due to
the fact that in these habitats there were only
three species for which the rate exceeded
1 m yr–1, whereas in other types of forests it
was much higher in many cases. In addition,
the overall species pool whose performance
in other types of forests can be compared is
very small, mostly because there are distinctive differences in habitat conditions between
pine woods and the forests for which the data
84
Anna Orczewska, Małgorzata Fernes
exist in the literature. Thus, as was mentioned
before, only a small proportion of species
common in both pine and deciduous forests
is available for comparisons. Migration rates
for all of the species from that set, even for
those which colonized pine woods reasonably quickly, when compared with other species in the pine woods studied here (D. carthusiana and M. trinervia) were lower than in
other habitats (Table 2). The exceptions were
E. sylvaticum when compared to recent alder
woods (Orcze wska 2009), and D. carthusiana, L. pilosa, M. trinervia, and M. muralis in
comparison with the studies by Dzwon ko
(2001b).
Despite the small pool of woodland species existing naturally in pine woods compared to the more fertile forest habitats,
some trends in the colonization capacity of
species can be described. In general, it can
be concluded that the pace of secondary
succession and migration of forest herbs in
such unproductive habitats proceeds more
slowly than in richer types of forest ecosystems, especially those with black alder. One
of the possible, ecological factors behind
this pattern is composition of the tree species of the recent stands. Black alder woods
with quickly decomposing leaf litter provide
better conditions for herb layer restoration
than pine stands with slowly decomposing
needles (Dzwon ko 2001a, b). Differences in
the species composition between ancient and
recent woods are still distinct even a few decades after afforestation. More studies in this
respect concentrating on the abiotic factors
that influence both the migration of woodland species and their successful establishment in the herb layer of recent pine woods
are required. Many authors, among others
F l i n n and Vel l e n d (2005) and Her my and
Verhe yen (2007), have also indicated such
a need. Thus, in order to answer the question
about which species from the ancient pine
woodlands are limited by dispersal mode and
which are recruitment-limited, more empirical data would be needed. For example,
experiments with the introduction of forest species into recent woods would help to
answer the questions about the mechanism
responsible for the observed distribution of
woodland species in recent woods. Seed sowing experiments and adult transplants are
among the group of surveys that would be
worth undertaking.
The studies in pine woods present distribution patterns of plants on a local scale. Thus,
due to the limited data set, the discussion is
relatively species-specific. Although some
trends in the colonization patterns can be observed, further research in this respect, based
on a bigger number of sites, is necessary.
ACKNOWLEDGEMENTS: We express
our gratitude to Artur Obidziński (Faculty of
Forestry, Warsaw Agricultural University) for his
comments on the previous version of the manuscript, to Małgorzata Scheiki-Bińkowska (University of Silesia) for preparing Figure 3, and to
Michele L. Simmons (English Language Centre,
University of Silesia, Katowice) for improving the
English style.
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Received after revision August 2010