Forest Ecology and Management 161 (2002) 109±122

Permeability of ancient forest edges for weedy

plant species invasion
Olivier Honnay*, Kris Verheyen, Martin Hermy
Laboratory for Forest, Nature and Landscape Research, University of Leuven, Belgium,
Vital Decosterstraat 102, B-3000 Leuven, Belgium
Received 24 May 1999; received in revised form 13 December 2000; accepted 15 January 2001

Abstract

Within highly fragmented forest landscapes, the ¯ux of matter, energy and species from the landscape matrix into the forest
fragment interior is an important ecological phenomenon. We studied the invasion of ancient forest edges by weedy plant
species, which are normally con®ned to the agricultural landscape matrix. We related this to edge orientation and to the
occurrence of natural and anthropogenical environmental edge gradients like microclimate and soil chemistry. The plant
community composition of the forest edge zone differed signi®cantly from the community composition of the forest core area,
and is characterized by a relatively high number of competitive and light demanding species. The maximal width of this edge
zone is more extensive at south facing (maximum ca. 20±23 m) than at north facing edges (ca. 0±3 m). However, most of
the weedy plant species were unable to penetrate the forest and were con®ned to the boundary between forest and arable land.
We found no negative edge effects on the more stress tolerant `true' forest species going further than 0±3 m (only at south
facing edges). The invasion pattern generally ®tted the edge gradient for photosynthetically active radiation (PAR). The
negative effects of fertilizer misplacement from adjacent arable land on forest soil chemistry (and community composition) are
also restricted to the boundary of the forest edge zone. Based on the derived penetration distances of the edge effects and on
the shape index of a forest patch it is possible to derive the minimal required forest patch area for forest plant species
conservation. # 2002 Elsevier Science B.V. All rights reserved.

Keywords: Forest invasion; Edge effects; Fertilizer drift; Forest microclimate

1. Introduction at the landscape scale is usually quanti®ed by edge

Forest fragmentation has led to increasing spatial
isolation and decreasing patch size of forest habitat
with subsequent negative implications for the survival
of the inhabiting natural populations (Harris, 1984).
An important aspect of the forest fragmentation
process is the relative increase of edge habitat, which
*
Corresponding author. Tel.: ‡32-16-32-97-73;
fax: ‡32-16-32-97-60.
E-mail address: olivier.honnay@agr.kuleuven.ac.be (O. Honnay).
density (McGarigal and Marks, 1995; Hargis et al.,
1998). Within high edge density landscapes, the ¯ux
of matter, energy and species from the landscape
matrix into the forest fragment interior and vice versa
is an important ecological phenomenon (Wiens, 1992;
Ryszkowski, 1992).
The main concern in a plant conservation context is
how the ¯uxes directed into the forest fragment may
in¯uence forest plant dynamics such as regeneration
and interspecies competition (Murcia, 1995). The
impact on forest plant dynamics and diversity will

0378-1127/02/$ ± see front matter # 2002 Elsevier Science B.V. All rights reserved.
PII: S 0 3 7 8 - 1 1 2 7 ( 0 1 ) 0 0 4 9 0 - X
110 O. Honnay et al. / Forest Ecology and Management 161 (2002) 109±122
depend on the penetration distance of the different
¯uxes, i.e. on the width of the edge zone. Edge width
can be de®ned as the part of the forest fragment where
environmental conditions differ signi®cantly from
those in the interior of the fragment, and, hence where
species composition and abundance also differ (Forman
and Moore, 1992). Edge width is the result of the
penetration distance of various environmental vari-
ables. A lot of the available studies, however, did not
directly measure these environmental edge gradients
and purely focused on the resulting vegetation patterns
(e.g. Ranney et al., 1981; Palik and Murphy, 1990;
Laurance, 1991; Chen et al., 1992; Fraver, 1994;
Kupfer, 1996; Medley, 1997; Buckley et al., 1997;
Esseen and Renhorn, 1998; Laurance et al., 1998;
Meiners and Pickett, 1999). The effects of naturally
occurring environmental edge gradients like soil
temperature, air temperature, photosynthetic active
radiation (PAR) and litter moisture on vegetation
patterns and, more speci®c, on alien plant species
invasion of the edge zone have also received attention
(e.g. Williams-Linera, 1990; Brothers and Spingarn,
1992; Matlack, 1993, 1994; Chen et al., 1995; Jose
et al., 1996; Williams-Linera et al., 1998). Depending
on edge orientation, age and physiognomy, the authors
report a transient zone up to maximal 50 m between
the landscape matrix and the typical forest core
microclimate. Brothers and Spingarn (1992) argued
the typical forest-core microclimate conditions to be a
relatively ef®cient protection against invasion of alien,
mostly weedy, species from the landscape matrix.
Almost all mentioned studies considered north or
south American forest edges. In the agricultural
landscapes of western Europe another important kind
of environmental variation across edges has been
underexplored. In addition to microclimate variation,
anthropogenic soil nutrient gradients may be expected
across forest edges as a result of intensive agricultural
activity in the surrounding landscape matrix. Two
different physical processes can generate such gra-
dients. First, forest edges are exposed to relatively
high atmospheric ion deposition rates because they
disturb the vertical wind pro®le and cause air turbulence
(Draaijers et al., 1988). At windward spruce forest edge
zones in the USA, Wheathers et al. (1995) noted on an
average three times more ion deposition (H‡, NH4‡,
NO3 , SO42 ) than in the interior of the forest. In a zone
of industrial livestock breeding in the northern part of
Belgium, De Schrijver et al. (1998) report a more than
two-fold increase of NH4‡ deposition at the edge of a
pine forest. Second a direct input of nutrients (mainly N,
P and K compounds) by drifting fertilizer application,
runoff and wind erosion from adjacent agricultural land
can be expected (Carlson et al., 1990; Kleijn and
Snoeijing, 1997).
It is known that arti®cial nutrient addition can have
major consequences for plant species diversity,
originating from changes in competitive relations
between species (Bobbink et al., 1998; Lee, 1998;
Turkington et al., 1998). In a forest environment this
means the competitive replacement of stress tolerant
forest plants species by more competitive species
(sensu Grime et al., 1988). For forest edge plants the
effects of atmospheric nitrogen deposition on wind
exposed ridges in southern Germany (Wilmanns et al.,
1986) and the effect of direct fertilizer misplacement
in Australia have been studied (Hester and Hobbs,
1992). These studies report an important increase of
the cover of competitive species in the forest edge
zone, and an increasing number of exotic species
suppressing the original forest ¯ora.
In this study we will investigate the effect of the
interaction between natural and arti®cial environ-
mental edge gradients on edge community composi-
tion and especially on forest edge invasibility by
weedy species which are usually con®ned to the
agricultural landscape matrix. We adopt a very broad
de®nition of invasion involving the expansion of a
species into a habitat where it is naturally not occurring
or only at very low importance values (see, in contrast,
the more restrictive de®nition in Thompson et al.
(1995)). We addressed the following questions:
1. Is the plant community composition of the forest
edge different from the forest core?
2. What is the effect of edge orientation on abiotic
edge gradients and on the penetration of weedy
plant species into the forest core?
3. Do increased nutrient levels in the edge zone
facilitate invasion or does forest microclimate
inhibits the invasion of non-forest species?
Our study area, located within the central Belgian
loam belt consists of large and ancient deciduous
forest fragments within a matrix of intensively used
arable land. The landscape is representative for many
western European agricultural landscapes.
 O. Honnay et al. / Forest Ecology and Management 161 (2002) 109±122 111

2. Materials and methods Table 1

Characteristics of the ®ve surveyed transects
2.1. Study area and data collection Transect Location Orientation Number of 3 m
 3 m quadrats
Our study area is situated in the central Belgian surveyed
loam belt, 25 km to the east of Brussels. It consists of A Bertem forest N 59
three large forest patches that are mainly deciduous B Grevens forest SSW 66
(Meerdaal forest (ca. 1200 ha), Bertem forest (ca. C Meerdaal forest SW 60
D Meerdaal forest NNE 70
200 ha) and Grevens forest (ca. 100 ha)). The forests
E Grevens forest SE 70
are classi®ed as Milio-Fagetum communities (sensu
Noirfalize, 1984). Within this study area we selected
®ve edges, at least 100 m long, and adjacent to
intensively used arable land (rotation of wheat and 20, 30, 40 and 50 m from the forest edge, and 10
sugar beets). All edges are at least 220 years old (i.e. repetitions at each distance, resulting in seventy
present on the oldest available topographical map), are 3 m  3 m quadrats per edge and an overall total of
very sharply delineated, and have a relatively homo- 350 quadrats (Fig. 1). Matlack (1990, 1994) used a
geneous and closed structure consisting of adventi- similar grid pattern. Ultimately 25 quadrats had to be
tious limbs of the boundary canopy trees and a shrub omitted because they were too disturbed (e.g. in the
belt under the canopy. Two of them are north facing vicinity of a ride). The location of the 0 m quadrat at
(N and NNE) and three are south facing (SE, SSE and the forest margin was determined by a visual analysis
SW) (Table 1). Study sites are covered with mature of the Ah soil horizon, indicating the boundary of
mixed deciduous forest (mainly Acer pseudoplatanus, (historical) plow activity.
Quercus robur, Fagus sylvatica and Prunus avium). In May 1998 the vegetation of each quadrat was
The same species are occurring in the shrub (0±2 m), surveyed using a decimal cover scale (Londo, 1974).
the coppice (5±10 m) and in the canopy layer (20±25 m). In September 1998 an additional survey was per-
Soils of the study sites are luvisols. formed to assess seasonal variation. Species not
Within each study site we established a regular present in the May survey were added to the species
53 m  30 m grid (i.e. a transect). Each transect versus quadrats data matrix. Only the herbaceous and
consisted of seven 3 m  3 m quadrats at 0, 5, 10, the shrub layer were surveyed. Coppice and canopy

Fig. 1. Each sampling grid (i.e. transect) consists of seven 3 m  3 m quadrats at 0, 5, 10, 20, 30, 40 and 50 m from the forest edge, together
with nine replicates at each distance, resulting in seventy 3 m  3 m plots per transect.
112 O. Honnay et al. / Forest Ecology and Management 161 (2002) 109±122
layers were omitted because they usually have been
planted. For each quadrat we derived the total plant
species richness and the number of species belonging
to two functional ecological species groups according
to Grime's CSR model (Grime et al., 1988; Hodgson
et al., 1995); we derived the number of competitive
species (i.e. competitives (C), competitive ruderals
(CR), C/CR, C/stress tolerant competitives (SC), CR/
CSR) and stress tolerant species (i.e. stress tolerators
(S), S/CSR) per quadrat together with their sum cover.
We also calculated the characteristic indicator value
for nitrogen (CIV-N), light (CIV-L) and reaction ®gure
(CIV-R) for each quadrat (Persson, 1981) based on the
Ellenberg indicator ®gures (Ellenberg et al., 1992).
Microclimate measurements (one transect each day)
were made between 11 and 18 September 1998, a
period with stable, clear weather. We only measured
between 12.00 and 14.00 h. In each quadrat we made
®ve measurements which were then averaged to one
value per quadrat. We also averaged ®ve measure-
ments in the open ®eld, at 50 m from the forest edge, to
one value. We measured soil temperature (8C), photo-
synthetically active radiation (PAR, 400±700 nm,
W/m2), air temperature (8C) and relative air humidity
(%), the latter three variables, all at 30 cm above
the ground level. Microclimate measurements were
expressed as fractions of the open ®eld values. For the
PAR we used a Solar Hog sensor (SDL 5000 series
solar cell PSU). All data were logged on line with the
Skye Datahog 2 (Skye Instruments Ltd., Powys, UK)
connected to a laptop computer.
In four of the ®ve transects (not in transect D), soil
was sampled to 5 cm depth (after litter removal) in
each quadrat (December 1998). For each distance,
samples were bulked together resulting in one bulk
sample per distance (i.e. seven per transect). The bulk
samples were analyzed according to the standard
methods of the Belgian Soil Service for available
cations (Ca2‡ and K‡) and available phosphate (both
Egner±Riehm method), carbon content (Walkley and
Black method), pH(KCl), available nitrogen (NO3 and
NH4‡) and total N-content (Kjeldahl±Lauro method),
see Hendrickx (1992) for details.
2.2. Analyses
Many studies dealing with the characterization of
edge effects suffered from pseudoreplication (sensu
Hurlbert, 1984). This means that many transects
(pseudoreplicates) were surveyed and analyzed, but
that they where all lying in the same forest edge
(Murcia, 1995). For a landscape scale study, proper
replicates should be located in different edges. To
avoid pseudoreplication, we averaged cover values,
CIVs, number of competitives/stress tolerators and
microclimate measurements per distance in each of
the ®ve studied transects. This resulted in ®ve repli-
cates per distance category, which are all based on the
average of 10 (vegetation data) or 50 (microclimate
data) measurements, and hence are considered to be
very reliable estimates.
Only in a ®rst data exploratory phase we used the
basic quadrat level data (n ˆ 325) in a correspondence
analysis (CA) to identify general patterns of plant
community variation. Vegetation data (% cover) were
square root transformed to lower the effects of extreme
values. CA-quadrat scores were then correlated with
transect orientation and quadrat distance to the forest
edge. We used a Mann±Whitney test and a Spearman
rank correlation, respectively (Siegel and Castellan,
1988).
We then focused on the penetration distance of the
¯ux of species, into the forest core. We applied a
generalized linear modeling (GLM) and analysis of
variance (ANOVA) procedure. Dependent variables
were species cover, CIVs and sum cover and number
of competitives and stress tolerators. Independent
model variables were distance to the edge (covariate),
edge orientation (N or S, ®xed factor) and the inter-
action term between distance and orientation. To
quantify penetration distances for dependent variables
exhibiting a signi®cant distance effect, we performed
Kruskal±Wallis pairwise comparisons between dis-
tances to specify the penetration distance (Siegel
and Castellan, 1988). For dependent variables with
a signi®cant GLM interaction term, penetration
distances were calculated for N and S exposed
transects separately. Note that for north facing edges
our observations are based on only two replicates per
distance (however, both based on 10 measurements).
Species only occurring in the 0 m edge quadrats (i.e.
unable to penetrate the forest edge zone any further)
were also omitted and were separately listed. The
same procedure was followed for the determination
of the edge width for the microclimate and the soil
variables. Microclimate variables were standardized
 O. Honnay et al. / Forest Ecology and Management 161 (2002) 109±122 113

by dividing them by the open ®eld value and
multiplying them by 100.
Besides spatial autocorrelation parallel to the forest
edge (resulting in pseudoreplication), that can be
avoided by a proper sampling design, autocorrelation
perpendicular to the forest edge is an intrinsic problem
of gradient studies. However, from scatter plots of
model residuals of neighboring quadrats it appeared
that the residuals were not spatially autocorrelated.
Hence, inferences from our model are valid. VAR-
IOWIN (Pannatier, 1996) was used to construct the
scatter plots.
For data analysis we used SPSS 9.0 (SPSS Inc.,
1999) and CANOCO for Windows 4.0 (Ter Braak and
Smilnauer, 1998). Plant species nomenclature follows
De Langhe et al. (1988).

Fig. 2. Quadrat scores of the CA (species cover values) on the ®rst two CA axes (pooled dataset, n ˆ 325). Transect identity and quadrat
distance to the forest edge are indicated, respectively.
114 O. Honnay et al. / Forest Ecology and Management 161 (2002) 109±122

3. Results Table 2
Correlation between CA quadrat scores and quadrat characteristics
(n ˆ 325)
The variation in community composition in the
surveyed quadrats can be attributed to the transect Quadrat distance Orientation (north or
orientation (CA axis I; eigenvalue 0.61) and to to forest edgea south facing transect)b
distance to the forest edge (CA axis II; eigenvalue CA x 0.07 11.32***
0.41) (Fig. 2, Table 2). CA y 0.69*** 1.96*
The ANOVA results (Table 3) con®rm the impor- a
Spearman correlation coef®cient.
tance of distance to edge and edge orientation in b
Mann±Whitney z-value.
determining community composition. More speci- *
0:01  P < 0:05.
***
®cally, the signi®cance of the interaction term P < 0:001.
`distance  orientation' for CIV-L, species number,
number and cover of competitives, and the cover of the 0±3 m quadrats (Table 4, Fig. 3). At south facing edges
invading species Galium aparine and Urtica dioica, U. dioica exhibits an increased cover up to 5±8 m, and
indicates that community composition and penetration G. aparine up to 10±13 m into the forest. There is an
distance strongly depends on transect orientation. increased cover of competitive species up to 20±23 m.
At northern edges signi®cantly increased values of Independent from edge orientation, Rubus fruticosus
mentioned response variables are restricted to the coll. has an increased cover value up to 10±13 m,

Table 3
F-values and signi®cance of the GLM ANOVA modela

Intercept Distance Orientation Distance  orientation Model

Species with decreasing cover values

Galium aparine 24.08*** 9.08** 24.08*** 9.08** 13.33***
Glechoma hederacea 5.73* 4.29* 5.74* 3.09 3.55*
Millium effusum 19.73*** 2.56 10.73** 0.88 6.85**
Poa trivialis 16.87** 9.08** 0.19 0.05 3.39*
Prunus serotina 5.15* 0.00 4.35* 0.13 3.76*
Ribes rubrum 5.41* 2.83 5.41* 2.83 3.31*
Rubus fruticosus coll. 40.63*** 11.18*** 1.47 0.22 4.85**
Sambucus nigra (seedling) 12.21** 4.08* 3.18 1.32 2.93*
Ulmus minor 4.75* 2.09 4.75* 2.09 2.74*
Urtica dioica 25.67*** 8.10** 17.56*** 4.64* 10.73***
Species with increasing cover values
Anemone nemorosa 10.5* 7.00* 3.6 4.5 18.1**
Convallaria majalis 0.52 7.70** 0.01 4.40* 5.50**
Community composition indicators
CIV-L 7.15* 9.76** 0.12 3.41* 2.71*
CIV-N 77.52 2.08 10.4* 0.13 2.39*
CIV-R 135.1 0.21 2.11 1.73 2.36*
Species number 230.0 4.39* 8.85** 3.68* 8.68*
Number of stress tolerators 78.94*** 0.13 10.00** 1.48 2.79*
Number of competitives 80.99*** 4.94* 3.23 4.20* 3.34*
Cover of stress tolerators 109.19*** 4.56** 45.71*** 1.91 7.12***
Cover of competitives 16.52*** 8.07** 8.01** 3.99* 6.36**
a
Response ˆ intercept ‡ distance ‡ orientation ‡ distance  orientation. Species cover values were square root transformed. Only
species with a signi®cant model F-value, and not occurring exclusively in the 0 m quadrats are presented.
*
0:01  P < 0:05.
**
0:001  P < 0:01.
***
P < 0:001.
 O. Honnay et al. / Forest Ecology and Management 161 (2002) 109±122 115

Table 4
Differences (P < 0:05) in mean cover, mean indicator value and mean species number between different distances to the forest edge according
to pairwise comparisons (indicated with letters)a

0±3 m 5±8 m 10±13 m 20±23 m 30±33 m 40±43 m 50±53 m

Species with decreasing cover values

Galium aparine (south) 0.86 a 0.82 a 0.31 ab 0.11 b 0.18 b 0.03 b 0b
Glechoma hederacea 4.41 a 2.17 a 0b 0b 0b 0b 0b
Hedera helix 1.44 a 0.61 ab 0.13 b 0b 0.05 b 0.03 b 0b
Moehringia trinervis 0.20 0.17 0.03 0 0 0 0
Poa trivialis 2.90 a 0.22 b 0b 0.07 b 0b 0b 0b
Rubus fruticosus coll. 13.00 a 5.82 b 2.14 b 0.64 c 0.44 c 1.07 c 0.94 c
Sambucus nigra (seedling) 0.24 0.39 0.13 0.18 0 0 0
Urtica dioica (south) 40.31 a 8.21 b 2.03 c 2.93 c 2.44 c 1.21 c 0.13 c
Species with increasing cover
Anemone nemorosa 1.28 16.43 48.24 37.60 33.48 36.87 34.16
Convallaria majalis (south) 0.0 1.47 6.10 0.40 0.20 4.40 3.87
Convallaria majalis (north) 0.20 0.80 4.05 11.75 4.85 22.25 10.45
Community composition indicators
CIV-L (south) 7.10 a 6.15 ba 5.85 b 5.15 c 5.12 c 4.40 c 4.24 c
CIV-L (north) 6.97 a 5.18 bc 5.35 bcd 5.42 bcd 4.96 b 5.1 bcd 4.84 d
Species number (south) 10.79 a 10.35 a 7.67 b 6.68 c 5.67 c 6.17 bc 6.38 c
Species number (north) 6.89 a 4.71 ab 5.9 ab 4.23 b 5.59 ab 4.83 ab 6.10 ab
Number of competitives (south) 9.09 a 5.69 b 2.10 b 1.11 c 1.0 c 1.03 c 0.93 c
Number of competitives (north) 2.42 a 0.85 b 1.05 b 1.18 b 1.88 b 1.28 b 1.10 b
Cover stress tolerators 1.82 a 20.78 b 55.50 c 43.74 bc 29.21 bc 55.59 c 48.79 bc
Cover competitives (south) 60.13 a 14.33 b 4.77 b 4.43 b 3.00 c 2.48 c 0.87 c
Cover competitives (north) 10.65 a 0.55 b 0.55 b 0.25 b 0.25 b 0.70 b 0.20 b
a
Only variables with a signi®cant distance effect in the full model are presented (see Table 3). Variables with a signi®cant interaction term
in the full model are evaluated for north and south facing edges separately when enough data available. Edge width, if derivable, is indicated in
bold.

Glechoma hederacea up to 5±8 m. Two species 4. Discussion

(Anemone nemorosa and Convallaria majalis) exhibit
a positive distance effect, their cover increases core
inwards. Table 5 lists the species which are exclusively
occurring in the 0±3 m quadrats, i.e. species which are
unable to penetrate the forest edge zone further than
0±3 m. The boundary of south facing edges is more
species rich than the one of north facing edges.
Also air and soil temperature vary with distance,
dependent on edge orientation (Table 6). There is no
temperature gradient at north facing edges while there
is an edge effect up to 20±23 m for air and 0±3 m for
soil temperature at south facing edges (Table 7, Fig. 4).
Edge width for PAR is 0±3 m (independent from edge
orientation) while there is no distance effect for
relative air humidity. Soil chemical characteristics
vary with distance independent from edge orientation.
For pH, Ca and K content, The Kruskal±Wallis
pairwise comparisons reveal a small edge zone.
4.1. Biotic edge gradients
The fringe of forest patches consists of a zone
where the plant community composition differs from
the community composition of the forest core area.
The studied edge communities are characterized by a
relatively high abundance of competitive and light
demanding species, especially at south facing edges.
Based on the sum cover of all competitive species, we
suggest an edge zone of at least 20±23 m at south
facing, and of 0±3 m at north facing edges. It is known
that the vegetation pattern with distance from the edge
varies with time since edge formation and also with
edge physiognomy (e.g. Matlack, 1993; Kupfer, 1996;
Esseen and Renhorn, 1998). In studies concerned with
late successional edges of temperate regions, like the
ones examined in this study, the edge widths reported
116 O. Honnay et al. / Forest Ecology and Management 161 (2002) 109±122

Fig. 3. Some typical examples of the biotic variation across the forest edge. The variation of the cover value of Rubus fruticosus coll. and
Urtica dioica (%) as a function of the distance to the forest edge is shown together with the sum cover (%) of all competitive plant species.
Bars represent 1 S.E.

by Palik and Murphy (1990) and Matlack (1994) in the
eastern part of the USA, are comparable with the ones
we found. Fraver (1994), studying mixed hardwood
forests in North Carolina (USA), mentions much
larger edge widths varying between 20 m (N) and
60 m (S) in at least 50 years old edges. However,
it is very dif®cult to assess and compare the edge
physiognomy and the corresponding degree of edge
closure in the different studies. Because edge physiog-
nomy affects the amount of light that reaches the
forest ¯oor and consequently also the intensity of
most other environmental edge gradients, it is a major
source of variation.
Although changes in community composition can
be detected up to 20±23 m in the forest (at south facing
edges), far most of the invading weedy species are
con®ned to the 0±3 m quadrats and only a very
restricted number of species is able to penetrate the
forest edge zone deeper than the established edge
boundary quadrats. These few deeper penetrating
species exhibit sharply decreasing cover values when
expanding into the forest core, results which are
corroborated by the ®ndings of Fraver (1994), Matlack
(1993) and Brothers and Spingarn (1992). Particularly
the latter authors explicitly studied forest edge
invasion by alien (mostly weedy) species in Indiana
 O. Honnay et al. / Forest Ecology and Management 161 (2002) 109±122 117

Table 5 species, although we found a negative effect on their

Species occurring exclusively at 0 m quadrats at south and north
sum cover value in the 0±3 m plots, both at south and
facing edges
north facing edges. This means that the edge effects on
Species occurring at south facing edges forest interior plant communities are relatively minor
Agrostis capillaris in our study area. Fraver (1994) also found no
Arctium spp.
Artemisia vulgaris
evidence of suppression of forest `interior' species,
Cirsium arvense even at the edge boundary. The question now remains
Deschampsia cespitosa which environmental factors prevent the invaders from
Festuca gigantea penetrating further into the forest edge and what is the
Holcus lanatus role of the soil chemistry gradients across the edge.
Heracleum sphondylium
Lamium album
Poa annua 4.2. Abiotic edge gradients
Poa nemoralis
Prunus avium (seedling) Although microclimate measurements were only
Ranunculus repens recorded during 1 week and one time period, and give
Rumex obtusifolius
Solanum dulcamara
no de®nitive representation of the micro climate at the
Stachys sylvatica site, our measurements are comparable with other
Stellaria media studies. Including diurnal microclimate variation,
Symphytum officinale Chen et al. (1995) already concluded that at temperate
Trifolium repens latitudes edge effect are strongest at SW facing edges
Taraxacum officinale
Vicia cracca
and weakest at NE facing edges. Our study con®rms
this for soil and air temperature in deciduous European
Species occurring at north facing edges
forests. At south facing, completely closed edges in
Agrostis capillaris
Holcus lanatus Pennsylvania and Delaware (USA), Matlack (1993)
Luzula multiflora could not detect microclimate gradients. At very
Poa annua recently established (<5 years), south facing edges,
Poa trivialis edge width ranged from 13 m (PAR) over 14 m (RH)
Stellaria media
up to 24 m for air temperature. Cadenasso et al.
Taraxacum officinale
Teucrium scorodonia (1997), studying NE and NW facing edges in New
Urtica dioica York (USA) report edge widths of 6.5±16.3 m for noon
temperature, of 8.4±17 m for soil temperature and of
11.8±25.1 m for relative air humidity. Although more
(USA) and report that most of the invaders at forest precise values were obtained in these studies, due to
margins have moved no more than two meter into the the used sampling design and the analysis methods,
forest interior. West and south facing edges appeared these ®gures are higher compared with our own
to be more rich in alien species. Meiners and Pickett results. This suggests, that ancient forest edges as the
(1999) report peak abundances of exotic species in ones studied have a relatively closed structure and are
New Jersey (USA) at 20 m of the forest edge. The relatively impermeable. This is due to the well
authors, however, studied relatively young, early developed shrub and coppice layer and to the
successional forests on former arable land. It is known sprouting of adventitious limbs of the boundary trees.
that soil disturbance, even historical, may increase the The reported pH values of the forest interior soil (ca.
likelihood of the invasion of a community (Hobbs and 3.5) correspond with the typical pH of the Ah and E
Huenneke, 1992). soil horizon under ancient forest developed on loess
The restricted invasive capacity of the weedy (Bossuyt et al., 1999). The increased pH values at the 0
species apparently prevents them to negatively affect and the 5 m quadrats suggest the input of fertilizer
the interior forest communities with their typical stress compounds from neighboring arable land (where the
tolerant forest plant species; we found no negative pH increases up to 6.5). It is known that liming and
effect of distance on the number of stress-tolerating fertilizing prevents soils under agricultural use from a
118 O. Honnay et al. / Forest Ecology and Management 161 (2002) 109±122

Table 6
F-values and signi®cance of the GLM ANOVA modela

Intercept Distance Orientation Distance  orientation Model

*** ** * **
Tair (8C) 5179.64 7.74 5.09 8.65 6.82***
Tsoil (8C) 761.91*** 5.16* 1.69 3.90* 3.83*
Relative humidity (%) 152.85*** 4.19 0.15 2.09 3.88*
PAR (W/m2) 84.16*** 8.00** 2.36 0.00 4.75**
pH(KCl) 463.14*** 10.08** 1.14 0.12 4.86*
Ca 113.87*** 24.52*** 1.20 0.69 8.83***
K 6.59*** 13.75** 0.42 0.41 6.59**
Mg 301.41*** 12.03** 5.08* 0.11 10.09***
P 29.87*** 6.48* 1.24 1.96 5.21*
Ntot 66.23*** 1.88 0.21 0.25 1.32
NO3 38.10*** 0.17 0.73 0.04 0.57
NH4‡ 62.33*** 4.23* 11.93** 4.02 4.84*
C 43.49*** 5.44 0.26 0.63 1.86
a
Response ˆ intercept ‡ distance ‡ orientation ‡ distance  orientation. Soil chemical characteristics in mg/100 g soil.
*
0:01  P < 0:05.
**
0:001  P < 0:01.
***
P < 0:001.

pH drop and soil degradation (pedological leaching)
which typically follows afforestation (HuÈttl and
Schaaf, 1995; Markewitz et al., 1998; Bossuyt et al.,
1999). Also Wilmanns et al. (1986) and Hester and
Hobbs (1992) noticed increased soil pH values at
forest edges which were exposed to drift of agro-
chemicals. The fertilizer misplacement hypothesis is
further supported by the higher K content of the
topsoil of the 0 m quadrats but is on the other hand not
re¯ected in an increased plant available P or increased
NO3 /NH4‡ levels. The N or P input from neighbor-
ing arable land is clearly not important enough to
cause persistent soil chemistry changes and to mask
the naturally occurring litter composition gradients in
forest soils, which re¯ect the canopy composition. We
cannot explain this, but probably this inconsistency is
related to the application form of the fertilizer. In the
study area, Ca and K are applied as powders and hence

Table 7
Differences in mean value of the environmental variables between different distances to the forest edge according to Kruskal±Wallis pairwise
comparisonsa

0±3 m 5±8 m 10±13 m 20±23 m 30±33 m 40±43 m 50±53 m

PAR 0.032 a 0.015 b 0.012 b 0.016 b 0.011 b 0.016 b 0.013 b

T (north) 94.38 94.39 94.48 94.38 94.36 94.34 94.39
T (south) 96.73 a 94.81 b 94.24 b 93.74 bc 94.02 c 93.79 c 93.35 c
Tsoil (north) 64.09 63.73 63.60 63.80 63.64 63.55 63.51
Tsoil (south) 68.61 a 64.42 ab 62.75 b 62.23 b 63.34 b 62.97 b 62.65 b
pH 4.62 a 4.10 a 3.58 ab 3.50 ab 3.45 ab 3.30 b 3.23 b
Ca 150.21 a 116.75 ab 89.75 b 70.00 b 56.75 b 53.00 b 43.33 b
K 17.00 a 14.25 ab 9.50 b 7.75 b 7.00 b 7.33 b 6.00 b
Mg 22.00 19.00 15.75 13.25 14.25 15.00 13.00
P 15.50 15.75 14.75 19.25 17.75 20.33 25.33
NH4‡ 2.76 3.09 5.68 3.36 2.84 3.55 1.88
a
Differences (P < 0:05) are indicated with letters. Only variables with a signi®cant distance effect in the full model are presented
(see Table 6). Variables with a signi®cant interaction term in the full model are evaluated for north and south facing edges separately.
The microclimate variables were standardized with respect to the open ®eld value (ˆ100).
 O. Honnay et al. / Forest Ecology and Management 161 (2002) 109±122 119

Fig. 4. Soil chemistry gradients (potassium and pH) and PAR variation across the forest edge. Bars represent 1 S.E.

are able to penetrate the forest edge boundary
(attached to soil particulates or by runoff of surface
water). P and N compounds to the contrary are applied
as granulates which ®nd more dif®culties to penetrate
the forest edge.
Finally, soil acidi®cation of the edge boundary
quadrats (which are expected as a result of the
oxidation of excessive NH4‡ deposition) could not be
noticed. In the Belgian loess belt these deposition rates
are rather low and, moreover, the NH4‡ atmospheric
deposition gradient across the forest edge may be too
¯at to be detected within a 50 m transect. De Schrijver
et al. (1998) have reported a gradient over more than
180 m in the sandy northern part of Belgium.
4.3. Linking biotic and abiotic gradients
We can conclude that most weedy species are
unable to penetrate the forest edge any further than the
boundary quadrats. This pattern very well ®ts the PAR
gradient. Increased PAR values are only noticed at the
0 m quadrats. Other authors have already suggested
that the penetration distance of weedy species is
mainly restricted because of the limited penetration of
light (Brothers and Spingarn, 1992). This seems to be
con®rmed for most of the encountered invasive
species, which are unable to penetrate the forest core
deeper than the 0 m quadrats. In contrast with mid-
successional species, the early successional species of
120 O. Honnay et al. / Forest Ecology and Management 161 (2002) 109±122
the landscape matrix have no adaptations for shade
tolerance (e.g. strong apical dominance). Hence, they
have no chance to penetrate and to reproduce under
canopy cover (Henry and Aarssen, 1997). The two
furthest penetrating species are R. fruticosus coll. and
U. dioica, both species which can grow under
relatively low PAR levels (`stress tolerating compe-
titives' sensu Grime). This pattern is also in accor-
dance with the patterns of increased nutrient levels
across the edge. The increased agro-nutrient input (Ca,
K) is restricted to the 0 m quadrats and the 5 m
quadrats (for pH) and works probably synergistically
with the increased PAR values of the edge. Generally
we can conclude, however, that the studied ancient
forest edges are relatively impermeable for weedy
species invasion.
4.4. Implications for conservation
Harrison and Bruna (1999) have recently suggested
that habitat fragmentation may be more a problem of
edge effects and less of spatial habitat con®guration
(i.e. patch isolation and patch area). For forest plant
species in forest fragments this has been con®rmed
by Honnay et al. (1999). Therefore, the importance
of the determination of the forest core area is a very
important conservation issue. The area of a forest
patch that is unaffected by edge effects not only
depends on the penetration distance of the edge
effect, and on the patch area but also on the shape of
the patch (Laurance and Yensen, 1991). The shape of a
habitat patch can be described by the Patton shape
index (PSI) which is de®ned by P/(200(pA))0.5 with P
the perimeter (m) and A the area (ha) of the forest
patch (Patton, 1975). Still according to Laurance and
Yensen, the core area can be calculated as follows:
total area …TA† affected area (AA) with AA ˆ
3:55d  PSI  …TA/10 000)0.5, where d (m) is the
penetration distance of the edge effects. Based on the
earlier derived penetration distances, it is possible to
assess the extent of edge effects for any given forest
fragment.
Acknowledgements
Thanks to Peter Cokelaere for help with data
preparation and data collection and the chemical
analysis of the soil samples and to Bea Bossuyt and
two anonymous reviewers for reading and discussing
the manuscript.
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