FLORISTIC ANALYSIS OF THE TWO WOODLAND–MEADOW ECOTONES DIFFERING IN ORIENTATION OF THE FOREST EDGE

Anna ORCZEWSKA, Aneta GLISTA

University of Silesia, Faculty of Biology and Environmental Protection Department of Ecology, Bankowa 9, 40-007 Katowice

e-mail: [email protected]

FLORISTIC ANALYSIS OF THE TWO WOODLAND–MEADOW ECOTONES DIFFERING IN ORIENTATION

OF THE FOREST EDGE

POLISH JOURNAL OF ECOLOGY(Pol. J. Ecol.)

53 3 365–382 2005

Regular research paper

ABSTRACT: Studies on the transition zones between plant communities using the statistical method by Matuszkiewicz, have been predomi-nantly conducted in forest communities or in ecotones between wetland and meadow com-munities. Although boundaries between forest and meadow (as a result of human pressure on the landscape, namely forest fragmentation) are of ecological interest, no previous attempts have been made to estimate the width of such ecotones using Matuszkiewicz’s approach. We applied the method in the studies on floristic changes across two, woodland-meadow ecotones (Arrhenathe-retum elatioris/Tilio cordatae-Carpinetum betuli typicum and Scirpetum sylvatici/Tilio cordatae-Carpinetum betuli corydaletosum communities), located at forest edges differing in their orien-tation. The aim of the research was to estimate the width and character of those ecotones and to investigate whether the possible differences in vegetation are related to the prevailing aspect (orientation) of the woodland edge.

To characterise the floristic composition of the two adjacent communities, the method of Greig-Smith’s square (16 quadrats of 4 m2 each) was chosen. To determine the floristic changes across each ecotone, transects 38 m in length and formed of 2 × 2 m quadrats were set perpendicu-larly to the woodland-meadow boundary. Diag-nostic species for the main communities were de-termined and their incidence along the transect was observed.

Differences in species composition that were observed along the transects allowed us to distin-guish three contrasting sectors. The first one rep-resents typical meadow communities, the second sector represents a typical ecotone, and the third one contains typical woodland communities.

Our results showed that the width of the eco-tones differed depending on the orientation of the woodland edge. In the transect between Ar-rhenatheretum elatioris and Tilio cordatae-Car-pinetum betuli typicum communities, situated on a SE-oriented forest edge, it was estimated at 10 m, whereas in the second transect, the tran-sition zone between the Scirpetum sylvatici and Tilio cordatae-Carpinetum betuli corydaletosum was narrower, reaching 6 m. This was located on a NW-oriented woodland edge. Our observations confirm the results of many other studies, which show that transition zones between woodland and meadow are wider on edges that receive more light, than in north-facing ones. The differences in the size of the transition zone at the two study sites are clear despite the fact that the two types of meadow communities differed in the number of mowing cycles per season. The Arrhenatheretum elatioris community is mown twice a year, and its ecotone with the woodland is wider than in the case of the Scirpetum sylvatici community. The latter is mown only once in the vegetation season but its transition zone remains 4 m narrower than in the first situation. It seemed that the increase in vegetation management (two mowing cycles

366 Anna Orczewska, Aneta Glista

instead of one) did not contribute to the reduced width of the ecotone.

When looking at the distribution of species representing different syntaxonomic groups, in both transects, no meadow plants from the Mo-linio-Arrhenatheretea class or other species of open habitats were recorded deeper than 6 m into the woodland interior. On the other hand, forest species from the Querco-Fagetea class were found much further into the area of meadow. Thus, no strong pressure of meadow communites on woods was recorded, but the opposite situation, where forest species extended into the meadow area, took place. Such a tendency was observed regardless of the aspect of the woodland edge.

Differences in physiological amplitude of species allowed us to distinguish, after R an-ney et al. (1981), a group of: meadow-oriented species, which neither occurred in ecotone nor woodland, strongly edge-oriented species, and strongly forest interior-oriented species, avoiding the ecotones and meadows. A group of species present along the whole transect (ubiquitous) was very poorly represented.

Contact zones between woodlands and meadows observed in the course of our studies and statistical analysis, give support to the model of a discontinuum.

In both ecotones studied, the edge effect was recorded; in transect 1 it can be described as a ‘double edge effect’ (variable = species richness, has both maximum and minimum in the ecotone close together); negative on the meadow side and positive on the woodland side of the ecotone, and at the second study site as the ‘positive edge effect’.

KEY WORDS: ecotone width, discontinuum, edge effect, ancient woodlands, southern Poland

1. INTRODUCTION

Two contrasting models that describe vegetation changes in the environment, the continuum and discontinuum concepts, pro-voked great discussion about community boudaries (van der Maarel 1976, Lorens 1990, Jeník 1992, and other references cited in those papers). Border zones between com-munities are also called contact or transition zones (Lorens 1990). The term most widely used to describe them, ecotone (gr. tonus = tension), was introduced by Clements (1905, see also van der Maarel 1976). It has been defined by him as “the line that connects the points of accumulated or abrupt change” (Harr is 1988). Thus, an ecotone

may be described as a tension belt (van der Maarel 1976). The typical feature of transi-tion zones between two communities is that the vegetation changes in one direction and is a combination of species occurring in the two adjoining communities (Fa l iński 1962, Traczyk 1960).

Floristic changes across transition zones between vegetation communities have been the object of many ecological studies (Tra-czyk 1960, Fa l iński 1962, Polakowska 1966, Matuszkiewicz 1972, Fa l ińska 1974, 1979a, b, R anney et al. 1981, Lorens 1984, Ba lcerkiewicz and Kasprowicz 1989, Lorens 1990, Kimsa 1991, L aur-ance 1991, Brothers and Spingarn 1992, Kwiatkowska and S ol ińska-Górnicka 1993, and others). In some cases the research has been undertaken in forest communities (Matuszkiewicz 1972, Kimsa 1974, Lo-rens 1984, 1990, Giec 1998). Forest-crop field boundaries (Dąbrowska-Prot 1995, Wójcik and Wasi łowska 1995, Choj-nacka-Ożga and Ożga 1999) and for-est–meadow ecotones (R anney et al. 1981, Ba lcerkiewicz and Kasprowicz 1989, L aurance 1991, Fraver 1994, Łuczaj and Sadowska 1997, L aurance et al. 1998, Honnay and Hermy 2000) have also been studied. These latter examples are of interest to ecologists as forest fragmenta-tion, a product of anthropogenic pressure on the landscape, has generated an increase in the amount of forest edges. As a result, un-desirable changes in the habitat of the for-est interior may take place (see Saunders et al. 1991, Harr is and Si lva-Lopez 1992, Murcia 1995).

One of the methods available for the study of ecotones is that proposed by Matuszkie-wicz (1972). It involves the determination of diagnostic species that make up the typical phytocoenoses in the dominant vegetation types, then analysis of vegetation changes along the ecotone based on the distribution of those species. Using formulas proposed by the author, statistical indices relating to spa-tial variation along the ecotone allow vege-tation changes to be described. The method also allows for the width of transition zones to be estimated.

Although several Polish ecologists have applied this method in their studies (Lorens

367Floristic changes across woodland-meadow ecotones

1984, 1990, Kimsa 1991, Giec 1998, Lo-rens 2001), it has not previously been used in research of forest-meadow boundaries. As the ecotones between those two entirely different ecosystems are of ecological inter-est, we focused our studies on the floristic changes in the herb layer along the wood-land-meadow gradients. The main aims of the research were:

– to describe changes in species compo-sition of herbaceous flora in the border be-tween meadow and woodland;

– to estimate the width and the character of the ecotones;

– to compare floristic changes in two transition zones differing in the orientation of the forest edge;

2. MATERIAL AND METHODS

Contact zones between meadow and woodland plant communities were compared in relation to the orientation of the woodland edge. General information about the study sites is presented in Table 1. The sites are lo-cated in southern Poland, on the central part of the Głubczyce Plateau (50o 21’ N; 17o 49’ E). They are located on two different sides of the same woodland. The woodland is an ancient one owned and managed by the state forestry service, whereas the meadows are in private ownership. At the first study site the meadow was mowed twice per year, and at the second site once. Both edges of the woodland are the same age and comparable in appearance.

The field and statistical methods used in course of the research followed, with slight modification, those described by Matusz-kiewicz (1972). Thus, in order to charac-terise the floristic composition of the two

contacting communities, the method of Greig-Smith’s square, with an area of 64 m2 divided into 16 quadrats of 4 m2 each, was used. Then transects, consisting of 19 2×2 m quadrats, with one end located in the typi-cal meadow community and the other in the typical woodland community, were used to study the herb layer vegetation through the contact zone between those two types of communities. Quadrats located in the meadow have a ‘–’ sign, whereas those origi-nating from the woodland are marked by a ‘+’ sign (Fig. 1.). Lists of species of vascular plants present in each of the quadrats, both in the typical communities and in the tran-sects, were recorded. The data were collected from early May to the middle of July in the year 2001. The nomenclature of plants fol-lows Mirek et al. (2002) and the names of vegetation communities follow those given by Matuszkiewicz (2001).

In the typical woodland and meadow community the frequency (f%) of each spe-cies was calculated.

f % = f

—16

·100%

(1)

The lists of species present in the typi-cal communities, and their frequencies, were prepared. Then, absolute differences in fre-quency D0 were calculated.

D0=| f1-f2 |(2)

Differences were tested for significance by comparing D0 with the critical value D0,01

Table 1. General information about the study sites.

No. Meadow association Woodland association Aspect of the woodland edge

Transect length

1. Arrhenatheretum elatioris Br.-Bl. ex. Scherr.1925

Tilio cordatae-Carpinetum betuli typicum Tracz. 1962

SE 38 m

2. Scirpetum sylvatici Ralski 1931 Tilio cordatae-Carpinetum betuli corydaletosum Tracz. 1962

NW 38 m


dcba

1

WOODLAND

3

2

01+

49+

8+

7+

6+

5+

4+

3+

2+

1+

1-BORDER

LINE

2-

3-

4-

5-

6-

7-

8-

9-

DCBA01- -11

MEADOW 2

3

2

1

4

8 m

m

2 m

2 m

2m

8m

D0,01= t0,01· 1— n1

1— n2

(3)

knowing that p*= f1+ f2—

2, q*=100 – p*;

where n1 and n2 are the numbers of samples that were used as the basis for the frequency calculations. The t0,01 value was

taken from the Student’s t test tables, for the 99% significance level, with v=n1+n2 de-grees of freedom. When D0>D0,01, the differ-ence is significant within 1% error. On this basis, species differentiating woodland and meadow communities were distinguished. Species for which the difference in the fre-quency in the two typical communities was not statistically significant (those occurring with similar frequency or only sporadically)

Fig. 1. Diagram of the research plots sited in the typical communities and in the transition zone between them.A, B, C, D – quadrats of the typical meadow community; a, b, c, d – quadrats of the typical woodland community; ‘-‘ – quadrats located in the meadow; ‘+’ – quadrats originating from the woodland.

2


were distinguished, too. In the groups of dif-ferentiating species for typical communities, those for which the frequency was higher in one of the communities (standard error ±5%), were included as well. Significant dif-ferences in their frequencies were confirmed by use of the non-parametric χ2 test (Siegel , Castel lan 2002), calculated using Statis-tica 5.0 software. For each of the 16 quadrats from the typical woodland (a) and meadow (b) community I value (‘informative value’) was calculated. The Ia index is the sum of D species regarded as locally differentiating for the woodland community, and Ib is the sum of D species regarded as differentiating for the meadow community.

Ia=ΣDa Ib=ΣDb (4)

Then, the W index, characterising the proportion of woodland and meadow ele-ment in each quadrat, was calculated accord-ing to the following formula:

Wa= Ia—

Ia+ Ib

Wb= Ib—

Ia+ Ib (5)

The I and W values calculated for each of the quadrats were used to determine the mean I and W values for the typical commu-nities: (Ia

–and Ib –) and (–Wb and –Wb).

In the next stage of the statistical analysis, the calculations of Ia and Ib, similar to those for the typical communities, were made for the transects. Thus the ‘relative value’ index (W*), defining the relative proportion of woodland or meadow community elements in a given quadrat of the transect, compared with the situation prevailing in the typical communities was calculated, according to the following formula:

Wa*= Ia—

(Ia+ Ib)· –Wb

Wb= Ia—

(Ia+ Ib)· –Wb

(6)Then, the ‘representativeness’ index (R),

indicating what percentage of the floristic components of the woodland or meadow communities occurs in a given quadrat of the transect, was calculated. According to Ma-

tuszkiewicz (1972), the ‘representative-ness’ index Ra is a ratio of Ia in a given square of the transect to the mean Ia of squares in the record of community a. Rb is the analogi-cal value for the community b.

Ra= Ia—Ia

·100% Rb= Ib—Ib

·100% (7)

Finally, two additional values were calcu-lated in order to help interpret the changes along the transects: SR, which is a sum of Ra and Rb, and QR, which is a value of absolute difference between Ra and Rb.

SR = Ra + Rb (8)QR = Ra – Rb (9)

The analysis of changes of W, R, S and Q along the transects allowed us to distinguish the ecotone within the transects.

The proportion of species of different syntaxonomic status (within each quadrat of the transects) was noted in order to provide additional information to help interpret the results when using the statistical approach of Matuszkiewicz (1972).

3. RESULTS

3.1. Transect 1. Arrhenatheretum elatioris–Tilio cordatae-Carpinetum betuli typicum Description of the typical communities

Within the Arrhenatheretum elatioris community 25 species were noted; of these 17 were regarded as locally differentiating. The number of species per quadrat ranged from 10 to 17, with 11 and 13 being the modus. In the Tilio cordatae-Carpinetum betuli typicum 27 species of vascular plants were noted, of which 20 were significantly more frequent in that community. The smallest number of species per quadrat was 8, the largest was 22.

3.1.1. Analysis of the transect based on the Matuszkiewicz approach

Sixty species were noted within the area of the transect, with 8 to 22 occurring per quadrat. The highest species richness was observed between the quadrats –2 and +3


(Fig. 2.). Meadow quadrats were poorer in species than those located in the woodland.

In the distal quadrats of the meadow community, first a gradual then, starting from the –4 quadrat, a rapid decrease in the number of species locally differentiating for the Arrhenatheretum elatioris community was noted (Fig. 3). At the same time, start-ing from quadrat –4, a rapid increase in the proportion of species differentiating for the Tilio cordatae–Carpinetum betuli typicum community was observed. Further into the woodland, the number of species of that group dropped slightly, then increased, but more slowly than in the earlier quadrats of the transect (Fig. 3).

Analysis of changes in the R, I, and W* indices allowed us to distinguish four sec-tions within the transect.

Sector I (quadrats –11 to –5) represents typical Arrhenatheretum elatioris communi-ty, where meadow species dominate (Fig. 3). The Ra, Ia and Wa* indices reach their highest values here, whereas Rb, Ib and Wb stay at zero level (Figs 4–6). In the latter case, exceptions were noted in quadrats –9 and –7, where they are different from zero. This is caused by the presence of Tilia cordata and Carpinus betulus, two species differentiating for the Tilio cordatae-Carpinetum betuli association. Species equally numerous in both communi-ties or present only sporadically within the transect, were noted.

Sector II (quadrats –4 to –3). The Ra, Ia and Wa* values drop quickly whereas Rb, Ib and Wb* indices increase rapidly (Figs 4–6). In the quadrat –3 the proportion of the spe-cies differentiating for the meadow commu-nity decreases. However, the proportion is still higher than that of the woodland species (Fig. 3).

Sector III (quadrats –2 to +1). The op-posite situation to that described in sector II takes place here. Thus, species differentiating for the woodland community reach a higher proportion than the meadow species (Fig. 3). The Rb, Ib and Wb* indices increase signifi-cantly, too (Figs 4–6). Species present only within the transect and not recorded in typi-cal meadow or woodland communities have their highest representation in this section of the transect.

Sector IV (quadrats +2 to +8). This part of the transect represents a typical Tilio cor-datae-Carpinetum betuli typicum commu-nity. The Rb, Ib and Wb* indices reach their highest values, whereas the Ra, Ia and Wa* values drop to zero (Figs 4–6).

3.1.2. Analysis of vegetation changes (proportion of species from different

syntaxonomic groups) along the transect

In the transect three zones, differing in the proportion of species representing differ-ent syntaxonomic groups, were detected as

02468

1012141618202224

-11 -10 -9 -8 -7 -6 -5 -4 -3 -2 -1 1 2 3 4 5 6 7 8

Quadrat number in the transect

Fig. 2. Number of species present in the quadrats (see Fig. 1) of transect 1 – Arrhenatheretum elatioris/Tilio cordatae-Carpinetum betuli typicum.


%

010203040

5060708090

100

-11 -10 -9 -8 -7 -6 -5 -4 -3 -2 -1 1 2 3 4 5 6 7 8

Quadrat number in the transectmeadow woodland

R

0

20

40

60

80

100

120

-11 -10 -9 -8 -7 -6 -5 -4 -3 -2 -1 1 2 3 4 5 6 7 8

Quadrat number in the transectRa Rb

I

0

200

400

600

800

1000

Quadrat number in the transectIa Ib

-11 -10 -9 -8 -7 -6 -5 -4 -3 -2 -1 1 2 3 4 5 6 7 8

Fig. 3. Proportion of locally differentiating species for transect 1 – Arrhenatheretum elatioris/Tilio cor-datae-Carpinetum betuli typicum (quadrat number – see Fig. 1).

Fig. 4. Spatial variation of the Ra/Rb indices of ‘representativeness’ (equation (7) – see text) for transect 1 – Arrhenatheretum elatioris/Tilio cordatae-Carpinetum betuli typicum (quadrat number – see Fig. 1).

Fig. 5. Spatial variation of the Ia/Ib indices of ‘informative value’ (equation (4) – see text) for transect 1 – Arrhenatheretum elatioris/Tilio cordatae-Carpinetum betuli typicum (quadrat number – see Fig. 1).


W*

0

0,2

0,4

0,6

0,8

1

1,2

Quadrat number in the transectWa* Wb*

-11 -10 -9 -8 -7 -6 -5 -4 -3 -2 -1 1 2 3 4 5 6 7 8

0%

10%

20%

30%

40%

50%

60%

70%

80%

90%

100%


Accompanying species

Rhamno-Prunetea

Phragmitetea

Artemisietea vulgaris

Agropyretea intermedio-repentis

Querco-Fagetea

Molinio-Arrhenatheretea


Fig. 6. Spatial variation of the Wa*/Wb* indices related to the proportion of woodland and meadow ele-ments in each quadrat (Fig. 1) (equation (6) – see text) for transect 1 – Arrhenatheretum elatioris/Tilio cordatae-Carpinetum betuli typicum.

Fig. 7. Changes in the proportion of species representing different syntaxonomic status occuring within transect 1 – Arrhenatheretum elatioris/Tilio cordatae-Carpinetum betuli typicum (quadrat number – see Fig. 1).

Fig. 8. Number of species present in the quadrats (see Fig. 1) of transect 2 – Scirpetum sylvatici/Tilio cordatae-Carpinetum betuli corydaletosum.


a result of the analysis of the vegetation data (Fig. 7).

Zone I (quadrats –11 to –5) is dominated by species characteristic of the Molinio-Ar-rhenatheretea class. Their proportionate occurrence is a few times higher than that of species of the other classes, that is from Querco-Fagetea and Agropyretea intermedio-repentis, and twice as great as the proportion of accompanying species.

Zone II (quadrats –4 to +1) includes sec-tors II and III that were distinguished as a result of the statistical analysis using Matusz-kiewicz’s approach. A significant decrease in the proportion of Molinio-Arrhenatheretea class species, accompanied by a doubling of the proportion of Querco-Fagetea class spe-cies, compared with that in the previous zone, is noted. The proportion of species characteristic of the Artemisietea vulgaris class is higher than in the previous zone. On the other hand, as from quadrat –3, represen-tatives of the Agropyretea intermedio-repentis class species are no longer present.

Zone III (quadrats +2 to +8). A high pro-portion of the Querco-Fagetea and Artemisie-tea vulgaris classes, and absence of Molinio-Arrhenatheretea class species is characteristic of this zone.

The analysis of the structure of the vege-tation and the calculations of the R, I and W* indices allowed us to distinguish a 10-metre wide transition zone between the Arrhenather-etum elatioris and Tilio cordatae-Carpinetum betuli typicum communities. It is located between quadrats –4 and +1 (Figs 3–6). A transition from meadow species to wood-land ones takes place in the ecotone (Fig. 7). The characteristic feature of that ecotone is a slight dominance of the woodland elements, reflected in particular in the spatial changes of I and R indexes along the transect.

3.2. Transect 2. Scirpetum sylvatici–Tilio cordatae-Carpinetum betuli corydaletosum

In the typical Scirpetum sylvatici com-munity 34 species were recorded; 25 of them were regarded as locally differentiating for the meadow community. In each quadrat, be-tween 15 and 23 species were noted, with 20 species occurring as the most frequent num-ber. The species richness of the Scirpetum

sylvatici community was three times greater than that of the woodland community. The woodland community was very poor in spe-cies, with only 14 noted in all the quadrats; 7 of them were locally differentiating for the Tilio cordatae-Carpinetum betuli corydaleto-sum association. The number of species per quadrat in that community ranged from 5 to 8, with 7 being the most frequent.

3.2.1. Analysis of the transect based on the Matuszkiewicz approach

In the Scirpetum sylvatici/Tilio cordatae-Carpinetum betuli corydaletosum transect 60 species of vascular plants were recorded. Species richness was several times greater at the end of the transect located in the meadow than in the quadrats situated at the opposite end, in the woodland (Fig. 8). The proportion of species locally differentiating for the meadow community is very high in the sector between quadrats –9 and –3; then drops slightly, reaching a proportion of less than 80%. Further into the woodland, the proportion of the meadow species decreases, to reaching zero in quadrats +3 and +4. No species differentiating for the Tilio cordatae-Carpinetum betuli corydaletosum community were observed in the sector between quadrats –9 and –2. A small proportion of the species from that group was noted in the area be-tween –1 and +2, whereas, starting from the +3 quadrat, their proportion increased sig-nificantly to reach an average level of 60% in quadrats representing the typical woodland community (Fig. 9).

As a result of the analysis of spatial variation of R, I and W* indices, three sec-tors within the transect between Scirpetum sylvatici and Tilio cordatae-Carpinetum betuli corydaletosum communities were recognised.

Sector I (quadrats –9 to –2). Initially the values of Ra and Ia indices stay at the same high level, then decrease quickly from quad-rat –5. However, the values of the Wa* index remain at a high level. The Rb, Ib and Wb* indices for the woodland community are at zero in that sector (Figs 10–12).

Sector II (quadrats –1 to +2). The sig-nificant decrease in the values of the Ra, Ia and Wa* indices is accompanied by a rapid


increase in the Rb, Ib and Wb* values. Sector II is the central part of the transect where 9 meadow species still occur and which is the initial place for the appearance of 6 wood-land species.

Sector III (quadrats +3 to +10). Minimal values for Ra, Ia and Wa* indices are noted, whereas the Rb and Wb* reach very high lev-els. The Ib index stays at a low level within the whole sector (Figs 10–12).

3.2.2. Analysis of vegetation changes (proportion of species from different

syntaxonomic groups) along the transect

The analysis of changes in the proportion of species representing different syntaxo-nomic groups along the length of the tran-

sect gave similar results to those obtained by Matuszkiewicz’s (1972) method and allowed us to distinguish three zones (Fig. 13).

Zone I (quadrats –9 to –2). Species from the Molinio-Arrhenatheretea class dominate here. The Phragmitetea class representa-tives are present along the whole sector. Although the species from the Artemisietea vulgaris and Querco-Fagetea classes occur there, their proportion is low. They start to play an important role in zone II (quadrats –1 to +2), where they are dominant groups. No representatives of the Phragmitetea class species are present beyond the end of zone II.

Zone III (quadrats +3 to +10) is domi-nated by species from the Artemisietea vul-garis and Querco-Fagetea classes. No species

Quadrat number in the transectmeadow woodland

Quadrat number in the transect Ra Rb

Fig. 9. Proportion of locally differentiating species for transect 2 – Scirpetum sylvatici/Tilio cordatae-Carpinetum betuli corydaletosum (quadrat number – see Fig. 1).

Fig. 10. Spatial variation of the Ra/Rb indices of ‘representativeness’ (equation (7) – see text) for transect 2 – Scirpetum sylvatici/Tilio cordatae-Carpinetum betuli corydaletosum (quadrat number – see Fig. 1).


from other syntaxonomic groups are repre-sented in this zone.

The analysis of changes of species com-position and of variation of R, I and W* in-dices allowed us to detect a 6-metre wide ecotone between the Scirpetum sylvatici and Tilio cordatae-Carpinetum betuli corydaleto-sum associations. It is located between quad-rats –1 and +2. The transition zone between those two communities is narrower than in the case of the Arrhenatheretum elatioris and Tilio cordatae-Carpinetum betuli typicum communities. Species typical for the two con-tacting communities are not mixed with the same abundance, but short sections charac-

terised by the dominance of either meadow or woodland species occur.

Changes in species composition along the border between meadow and woodland reflect changes in habitat conditions along that microclimatic gradient. Differences in physiological amplitude of species allowed us to distinguish, after R anney et al. (1981), groups of plants predominantly or exclusively occurring in meadow, ecotone or forest inte-rior zones. A group of meadow-oriented spe-cies, which neither occurred in ecotone nor woodland was represented for example by: Cerastium holosteoides, Poa pratensis, Rumex acetosa, Veronica arvensis, Bromus hordea-

Quadrat number in the transectIa Ib

W*

Quadrat number in the transectW*a W*b

Fig. 11. Spatial variation of the Ia/Ib indices of ‘informative value’ (equation (4) – see text) for transect 2 – Scirpetum sylvatici/Tilio cordatae-Carpinetum betuli corydaletosum (quadrat number – see Fig. 1).

Fig. 12. Spatial variation of the Wa*/Wb* indices related to the proportion of woodland and meadow elements in each quadrat (Fig. 1) (equation (6) – see text) for transect 2 – Scirpetum sylvatici/Tilio cor-datae-Carpinetum betuli corydaletosum.


ceus, Trifolium pratense, Carex acutiformis, Carex brizoides, Carex hirta, Glyceria maxi-ma and Holcus lanatus. Among the strongly edge-oriented species there were: Lythrum salicaria, Caltha palustris, Rumex sanguin-eus, Veronica beccabunga, Scrophularia nodosa and a few others noted only in one quadrat. The following plants were classified as strongly forest interior-oriented, avoiding the ecotones and meadows: Melandrium ru-brum, Moehringia trinervia, Carex sylvatica and Pulmonaria obscura. These last three species have the status of ancient woodland indicator species (sensu Peterken 1974) for Poland (Dzwonko and Loster 2001). Species present along the whole transect (ubi-quitous) were restricted to Glechoma hedera-cea in transect 1 and Acer pseudoplatanus in transect 2.

4. DISCUSSION

Using Matuszkiewicz’s method in the current study, changes in the vegetation along the forest–meadow ecotones were described. Differences in species composi-tion that were observed along the transects allowed us to distinguish three contrasting sectors. The first sector represents typi-cal meadow communities. This is charac-terised by high values of Ia, Wa, Ra and by the high proportion of locally differentiat-ing species for the meadow community (Figs 3–6, 9–12). The second sector repre-

sents a typical ecotone (following analy-sis of species from different syntaxonomic groups, it was shown to consist sometimes of two shorter sectors). An equilibrium in the proportion of species differentiating for meadow and for forest communities is reached in this sector, and the values of I, W and R indices for the two communities are comparable (Figs 3–6, 9–12). The width of the ecotone in this study is either 6 or 10 m. The third sector contains typical wood-land communities. The values of Ib, Wb, Rb and the proportion of locally differentiating species are characteristic for these woodland communities (Figs 3–6, 9–12). Successive changes in the proportion of species repre-senting different syntaxonomic groups are observed within the area of the transects. The widths of the sectors estimated on that basis overlap with those indicated by use of the Matuszkiewicz method.

The results show that the transition from typical meadow community to typical forest community takes places within an area of 6 or 10 m. These results are in general accor-dance with those obtained by R anney et al. (1981), Honnay and Hermy (2000) and Sławski (2001), who estimated the width of forest-meadow boundaries to be 15 m, 5–10 m, and 5–20 m respectively. However, it should be noted that because of the differ-ent methods used by these authors only an approximate comparison of the results can be made.


accompanyingspeciesPhragmitetea

ArtemisieteavulgarisAlnetea glutinosae

Querco-Fagetea

Molinio-Arrhenatheretea

Fig. 13. Changes in the proportion of species representing different syntaxonomic status occuring with-in transect 2 – Scirpetum sylvatici/Tilio cordatae-Carpinetum betuli corydaletosum (quadrat number – see Fig. 1).


Studies conducted on the ecotones be-tween forest communities show that the width of that zone is greater than that be-tween non-forest communities. One of the first studies of that phenomenon, made by Traczyk (1960), estimated the boundary to be from 5 to 15 m wide, depending on the floristic similarity of the communities (Cariceto elongatae-Alnetum chrysospleni-etosum – Querceto-Pinetum – to 5 m; Cari-ceto elongatae-Alnetum – Sphagnetum medii pinetosum – to 10 m; Cariceto elongatae-Alnetum chrysosplenietosum – Sphagnetum medii pinetosum – 10-15 m). The width of the ecotone between Querco-Carpinetum medioeuropaeum and Circaeo-Alnetum com-munities was estimated at 20-30 m (Pola-kowska 1966). However, the results were obtained by methods different from those used by Matuszkiewicz (1972), who calculated that the contact zone between Potentillo albae-Quercetum and Querco-Carpinetum was 60 m. Similar results were obtained by Lorens (1990), who calculated the widths of the ecotones for the following pairs of communities: Dentario glandulo-sae-Fagetum – Tilio-Carpinetum – (66 m); Dentario glandulosae-Fagetum – Abietetum polonicum – (70 m); Ledo-Sphagnetum mag-ellanici – Vaccinio uliginosi-Pinetum – (22 m); Vaccinio uliginosi-Pinetum – Leuco-bryo-Pinetum – (36 m); Peucedano-Pinetum – Querco-Piceetum – (12 m); Abietetum po-lonicum – Leucobryo-Pinetum – (18 m). The ecotone width ranged from 12 to 70 m, de-pending on the degree of floristic similar-ity in the contacting communities. Kimsa (1991) estimated the boundary between Abietetum polonicum and Dentario glandulo-sae-Fagetum communities as 16 m, whereas studies by Giec (1998) found the contact zones between different types of forest com-munities (Dentario enneaphyllidis-Fagetum, Tilio-Carpinetum, Luzulo pilosae-Fagetum, Calamagrostio-Quercetum, Pino-Quercetum, Querco roboris-Pinetum, Peucedano-Pine-tum, and Leucobryo-Pinetum) being much shorter (2–16 m), compared with other re-sults from forest communities. Research on ecotones between associations representing different types of wetland and meadow com-munities (Lorens 2001) estimated widths of 1–17 m, with 3 m being the most frequent.

Results from the above authors show that generally there is a greater variation in the widths of ecotones between forest com-munities than between non-forest ones. Al-though in some cases forest contact zones were of comparable size to meadow–wetland ecotones (Lorens 2001) or to the woodland–meadow boundaries present in this study, they were usually much wider.

Many authors (Traczyk 1960, Pola-kowska 1966, Matuszkiewicz 1972, Lo-rens 1984, 1990, Kimsa 1991, Giec 1998, Lorens 2001) have shown that boundaries between communities which are floristically similar are not sharp and that the transition of species composition takes place across a wide area, whereas communities differing in floristic composition have much narrower and sharper contact zones. Such a relation-ship between the size of the transition zone and the degree of floristic similarity between the two contacting communities was ob-served in our study. Meadow and woodland, as two very contrasting ecosystems, had nar-rower transition zones than most of the other types of communities. However, although these might be expected to be only a few metres wide, in one case the transition zone between meadow and woodland extended to 10 m. Other factors (discussed later) could influence that situation. However, in com-parison with the results obtained by Ma-tuszkiewicz (1972), Lorens (1984, 1990), Kimsa (1991), Giec (1998), and Lorens (2001), who applied Matuszkiewicz’s method and defined the character of the transition zones studied as a relative discontinnum (=although one phytocoenosis changes into the second one in a continuous way, there are some discontinuities in the transition zone – see Matuszkiewicz 1972), contact zones between woodlands and meadows ob-served in course of our studies and statistical analysis, more support the model of a discon-tinuum. Figures 14 and 15, showing the spa-tial variation of the Q index, support these findings.

Our studies showed that the width and the character of the ecotones differed de-pending on the orientation of the woodland edge. The first transect, between Arrhenather-etum elatioris and Tilio cordatae-Carpinetum betuli typicum communities, was situated on


a SE-oriented forest edge. The width of its ecotone was estimated at 10 m. In the sec-ond transect the transition zone between the Scirpetum sylvatici and Tilio cordatae-Car-pinetum betuli corydaletosum was narrower and reached 6 m. This was located on a NW-oriented woodland edge.

It has often been observed that ecotones at south- and south-west facing edges tend to be wider than those at north-facing ones (Yahner 1988, Fraver 1994, Murcia 1995, Ferr is and Car ter 2000, Honnay and Hermy 2000). This is due to the effects of insolation and of the prevailing wind (Fer-r is and Car ter 2000). Fraver (1994) stated that edge effect influenced south-facing for-

ests to a depth of 60 m into the stand, where-as on north-facing edges the effect was ap-parent only 10–20 m into the forest interior. According to Honnay and Hermy (2000) the widths of ecotones on edges oriented to the south reached 10 m, whereas on north-facing edges they were only 0–5 m. Honnay and Hermy emphasised that in their study sites environmental gradients at north facing edges were much steeper, so the conditions typical for the forest interior were reached within a very short distance from the edge.

Only Sławski (2001) has obtained dif-ferent results. His studies showed that transi-tion zones between woodland and meadow were wider in edges which received less light,

Sr Qr

Quadrat number in the transectSR QR

Sr Qr

Quadrat number in the transectSR QR

Fig. 14. Spatial variation of the Qr and Sr indices (equations (9) and (8) – see text) for transect 1 – Arrhenatheretum elatioris/Tilio cordatae-Carpinetum betuli typicum (quadrat number – see Fig. 1).

Fig. 15. Spatial variation of the Qr and Sr indices (equations (9) and (8) – see text) for transect 2 – Scirpetum sylvatici/Tilio cordatae-Carpinetum betuli corydaletosum (quadrat number – see Fig. 1).


that is those with west and north aspects, than in well lit, south-facing edges. He described the migration of species from open spaces into the forest interior, which was widely observed at south-facing edges, whereas at north-facing ones there was an expansion of forest species in the opposite direction. Ac-cording to Sławski these observations show that at south-facing edges there is stronger pressure from non-forest communities on forests than at edges oriented to the north. This might be due to rapid changes in the microclimatic conditions at southern edges (Young and Mitchel l 1994, Wójcik and Wasi łowska 1995).

No similar pattern in the distribution of plants was observed in our studies. In tran-sect 2, which was located on the NW edge of a woodland, the ecotone zone penetrated the wood 2 m deeper than in transect 1, located on the south-facing edge. Besides, at this lat-ter site the transition zone started 6 m earlier within the meadow area than at the site with the north-facing edge (Figs 3–6, 9–12). Thus, in the case of the first transect, the ecotone was 4 m wider than at the second study site but started earlier on the meadow side of the boundary.

When looking at the distribution of spe-cies representing different syntaxonomic groups, in both transects no meadow plants from the Molinio-Arrhenatheretea class or other species of open habitats (Phragmitetea class) were recorded deeper than 6 m into the woodland interior. On the other hand, forest species from the Querco-Fagetea class were found much further into the area of meadow (Figs 7, 13). Thus, no strong pressure of mead-ow communites on woods was recorded, but the opposite situation, where forest species extended into the meadow area, took place. Such a tendency was observed regardless of the aspect of the woodland edge. Light de-manding meadow species cannot survive in shaded conditions, thus such a pattern in their distribution is not surprising. Besides, the woodland edges at both study sites had well developed shrub vegetation. This would also contribute to the rapid decrease in the number of meadow species in the woodland part of the transects. This is in accordance with the results obtained by Brothers and Spingarn (1992) who studied the behav-

iour of alien species along the edges of for-est fragments. Although alien species were present at forest edges, their richness and frequency dropped sharply inside the forests. Thus, forest interiors were relatively free of alien species mainly because of the develop-ment of a dense wall of bordering vegetation, contributing to the reduction of the amount of light and wind speed.

The differences in the size of the transi-tion zone at the two study sites are clear de-spite the fact that the two types of meadow communities differed in the number of mowing cycles per season. Even though the Arrhenatheretum elatioris community is mowed twice per year, its ecotone with the woodland is wider than in the case of the Scirpetum sylvatici association. The latter is mowed only once in the vegetation season but its transition zone remains 4 m narrower than in the first situation. These results allow as to conclude that although, as Wójcik and Wasi łowska (1995) state, mowing may be a strong factor contributing to the narrower transition zones between meadows and for-ests, it is not the factor of greatest impor-tance. In some cases the character and width of ecotones is predominantly dependent on the orientation of the woodland edge. It seemed that in these situations the increase in vegetation management (two mowing cy-cles instead of one) did not contribute to the reduced width of the ecotone.

The increase in species richness and abundance in the ecotones, compared with that observed in the two main communities, ascribed to edge effect, is often noted in the literature (Odum 1963, Kershaw 1978, Fa l ińska 1991). Although, as Łuczaj and Sadowska (1997) mention, the idea of the edge effect has been criticised, it has been recorded in many studies (Matuszkiewicz 1972, Ba lcerkiewicz and Kasprowicz 1989, Łuczaj and Sadowska 1997, Lo-rens 2001), including ours. According to the terminology proposed by Łuczaj and Sa-dowska (1997), the edge effect recorded in transect 1 can be described as a ‘double edge effect’ (variable = species richness, has both maximum and minimum in the ecotone close together); negative on the meadow side and positive on the woodland side of the ecotone (Fig. 2). At the second study site there was


a tendency for species richness to decrease from the meadow towards the woodland in-terior. However, within the ecotone the num-ber of species increased. Thus, the ‘positive edge effect’ is observed there (Fig. 8). Figures 14 and 15 confirm the presence of edge effect at both study sites. Our results are different from those obtained by Łuczaj and Sad-owska (1997), who observed only a very weak edge effect in herbaceous plant rich-ness along the boundary between forest and meadow.

5. CONCLUSIONS

• The width of woodland-meadow eco-tones was in general narrower in comparison with studies undertaken by other authors who compared different combinations of vegetation communities.

• Although the method that we applied did not take the abundance of species into consideration, only the species composition, it was appropriate for the analysis of vegeta-tion changes along the boundaries between woodland and meadow. It allowed us to clas-sify the border as discontinuum.

• The differences in the character of the plant communities and in the width of the transition zone between the two pairs of communities studied seem to be influenced by the orientation of the woodland edge.

• The presence of an edge effect (double or positive) was detected in the study sites.

• Transition zones between meadow and woodland are ecologically interesting. Re-lated aspects such as the character and rate of changes of forest–meadow boundaries af-ter the cessation of mowing, or the changes in width of an ecotone in relation to pheno-logical changes within two communities, are worthy of future study.

ACKNOWLEDGEMENTS: We are very thankful to Dr John Parker † (formerly with the Forestry Commission, UK) for his kind help in improving the language of the paper.

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(Received after revising May 2005)

FLORISTIC ANALYSIS OF THE TWO WOODLAND–MEADOW ECOTONES DIFFERING IN ORIENTATION OF THE FOREST EDGE

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