Landscape Ecology 18: 253263, 2003.

2003 Kluwer Academic Publishers. Printed in the Netherlands.

253
Research article
Arthropod reaction to landscape and habitat features in agricultural
landscapes
Ph. Jeanneret
1
, B. Schpbach
1
, L. Pffner
2
& Th. Walter
1
1
Swiss Federal Research Station for Agroecology and Agriculture (FAL), Reckenholzstrasse 191, 8046 Zurich,
Switzerland
2
Research Institute of organic agriculture, Ackerstrasse, Postfach, 5070 Frick, Switzerland
E-mail to: philippe.jeanneret@fal.admin.ch
Key words: arthropods, biodiversity, canonical correspondence analysis, environmental control, landscape and
habitat inuence, variation partitioning
Abstract
Determining explanatory environmental factors that lead to patterns of biodiversity in cultivated landscapes is an
important step for the assessment of the impact of landscape changes. In the context of an assessment of the effect
of agricultural national extensication programme on biodiversity, eld data of 2 regions were collected according
to a stratied sampling method. A distribution model of 3 indicator species taxa (butteries, spiders, and carabid
beetles) is related to inuencing factors by means of multivariate statistics (CCA, partial CCA). Hypothetical
inuencing factors are categorised as follows: (1) habitat (habitat type, management techniques) and (2) landscape
(habitat heterogeneity, variability, diversity, proportion of natural and semi-natural areas). The correlation models
developed for spider, carabid beetle and buttery assemblages revealed that there are no general rules relating
species diversity to habitat and landscape features. The relationship strongly depends on the organism and on the
region under study. Therefore, biodiversity response to landscape and habitat changes has to be identied by means
of a multi-indicator concept in different landscape situations.
Introduction
The structure of a given biotic community is gener-
ally related to two classical models: the environmental
control model (e.g., Whittaker 1956) and the biotic
control model (e.g., Southwood 1987). These two
groups of inuencing factors are not mutually exclu-
sive, but complementary together with other factors
like historical events and disturbances of various kinds
(Quinn and Dunham 1983). On a broader scale, land-
scape characteristics are relevant explanatory variables
for plant and animal communities because they dene
the ecosystem arrangement and interactions (Forman
and Godron 1986; Forman 1995) and thus affect pop-
ulations via complementation and supplementation
processes (Dunning et al. 1992). The spatial arrange-
ment of habitat elements and the spatio-temporal het-
erogeneity of the landscape are essential for species
diversity (Burel 1992; Huston 1995). In the agricul-
tural landscapes in particular, undisturbed habitats,
their proportion to cultivated elds and their position
in the landscape, play an important role as refuges
and sources of individuals for recolonization (Den-
nis and Fry 1992; Lys and Nentwig 1994; Pffner
and Luka 2000). However, it is difcult to gener-
alise as to the signicance of spatial structure because
species ecology and dispersal abilities are different for
every organism (Burel and Baudry 1995; Burel and
Baudry 1999). For many arthropods, survival in agri-
cultural landscapes depends on the suitability of the
habitats, which is largely inuenced by eld manage-
ment, but also on the characteristics of the surrounding
landscape.
254
Our study aims to analyse, discriminate and com-
pare with correlative models two components of the
environmental control, i.e., the habitat and the land-
scape, acting on three arthropod taxa, i.e., spiders,
carabid beetles and butteries, collected and observed
in two regions that are different as to their landscape
congurations and land uses. The results of both re-
gions were discussed separately in Jeanneret et al.
(2002, and 2002 submitted), and showed that the three
arthropod taxa react differently to the habitat and land-
scape features in both regions. In this paper, we will
compare the reaction of the three arthropod commu-
nities between the two regions. We hypothesized that
one given taxa would react differently to the habitat
and landscape features depending on the region.
Both control components are divided into poten-
tial explanatory factors: (1) habitat descriptors: the
plant species richness, the habitat type; and (2) land-
scape descriptors: the surrounding habitat variability
and heterogeneity, an index of landscape pattern, the
surrounding land use.
As a basic habitat characteristic, plant species
richness is supposed to have a major inuence on
the spider, carabid beetle and buttery assemblages.
For spiders and carabid beetles, higher plant species
richness offers more diverse habitat structure (archi-
tecture) and more niches for prey (Strong et al. 1984).
Higher plant species richness represents more host
and feeding plants in time and space that should in-
uence buttery assemblages (Erhardt 1985; Sparks
and Parish 1995). In our context, the habitat type
distinguishes crops (cereal elds and high intensity
meadows) from ecological compensation areas (ECA)
and forest edges. ECA are elds set aside encom-
passing traditional landscape elements as well as new
types of biotopes which are designed to enrich the
agricultural landscape (Herzog et al. 2001). The habi-
tat type should play an important role in determining
the species composition of the arthropod taxa under
study as it represents the sum of the abiotic factors
characterising the sites. On the landscape scale, vari-
ability and heterogeneity of the surrounding habitats
may inuence the biodiversity measured at a given
point within the landscape (Duelli 1997). To comple-
ment these simple measurements of spatial pattern, the
D1 index of dominance based on information theory
was used (ONeil et al. 1988).
Nevertheless, the causes explaining the species dis-
tribution at landscape scale are usually very diverse
and habitat and landscape descriptors as proposed in
this paper are not supposed to be able to explain all
aspects of this distribution. Therefore, we also intro-
duced the spatial position of the sampling sites (rep-
resented by geographical coordinates) since it can be
considered as evidence for the various processes that
have generated the species distribution. In this study
the detection of spatial variation was not analysed per
se but the spatial position of the sites was integrated
as explanatory factor for playing the role of a syn-
thetic indirect descriptor of the unmeasured factors
as dened by Borcard et al. (1992) and Borcard and
Legendre (1994).
Method
Regions, sampling methods
The study was carried out in 2 regions of the cen-
tral Swiss Plateau: region 1 (Ruswil, 20 km NW of
Lucerne) and region 2 (Rafz, 20 km NW of Zurich).
Region 1 has undulating hills situated between an alti-
tude of 650 and 800 m. It comprises a total surface of
885 hectares, mainly consisting of arable land (15%),
grassland (59%), and forests (17%). Four ECA habi-
tat types, usually small areas of approx. 400 m
2
, can
be found in the perimeter, namely extensively used
meadows (no fertilisation, late mowing), low inten-
sity meadows (restricted fertilisation, late mowing),
hedgerows and standard fruit trees in traditional or-
chards. Region 2 has a at relief and is situated at a
mean altitude of 450 m. It comprises a total surface of
1016 hectares, mainly consisting of arable land (47%),
grassland (5%), forests (20%), gravel pits (11%), and
special cultures (6%). In region 2, the same ECA types
occur except that wild ower strips replace standard
fruit trees. The difference between both land uses is
the proportion of arable land and grassland, and the
presence of gravel pits and special cultures in region 2.
Spiders, carabid beetles and butteries were
recorded according to a stratied sampling method.
ECA, cultivated areas and forest edges were dened
as strata. The number of samples per ECA type was
determined in proportion to the number of elements
in each type occurring in the study areas (Table 1).
This attribution of ECA samples was possible because
the size of the elements in the ECA types was very
similar. The minimum number of samples was given
by the ECA type having the smallest number of el-
ements, i.e., hedgerows, and the number of samples
of the other ECA types was calculated proportionnally
to it. Seven highly intensive meadows and 20 winter
255
wheat elds in region 1 and region 2, respectively,
were chosen to serve as references for the cultivated
areas because they are predominant in the landscape.
Seventeen observation sites were set up along the for-
est edge belonging to 3 forest plots in region 1 and 6
sites along the forest of 2 plots in region 2.
Spiders and carabid beetles were collected in 1997
and butteries observed in 1998 in the 58 (region 1)
and 51 (region 2) sites. Spiders and carabid beetles
were collected with 3 pitfall traps per site, during
5 weeks (during the rst 3 weeks of May and last
2 weeks of June), as proposed by Duelli (1997) to
optimise the number of species caught compared to
the sampling effort (see also Section Methodological
aspects). This relatively short sampling period is nec-
essary to ensure uniformity of the habitat conditions
in winter wheat elds and high intensity meadows. A
longer sampling period would include a habitat change
(i.e., winter wheat - harvest mid July - new crop, e.g.,
winter barley). This change would strongly affect the
comparison between habitat types. The pitfall traps
used in this study consisted of funnel traps of 10.5 cm
in diameter containing 2 cm of 90% alcohol/water so-
lution. The 3 pitfall traps, and 4 (region 1) and 5 weeks
of sampling per site are pooled for the analysis. Due to
bad weather conditions in region 1, spiders and cara-
bid beetles were identied and analysed for 4 weeks
only. Butteries were observed across a 0.25 ha area,
5 times for 10 min each from the beginning of May to
the end of August. Butteries were monitored between
10.00 a.m and 5.30 p.m in sunny weather conditions,
with no or light wind and a minimum temperature of
18

C. At forest edges, butteries were recorded along

the edge. The 5 observation runs per site are pooled for
the analysis. At each of the observation sites, the veg-
etation was assessed over an area of 100 m
2
according
to the Braun-Blanquet method.
Measurement of environmental control
The explanatory environmental variables are divided
in three sets of descriptors (Table 2): (1) the habitat;
(2) the landscape; and (3) the space.
Habitat descriptors. 1. Plant species richness: num-
ber of plant species on 100 m
2
. 2. Habitat types:
habitats were assigned to the 8 types listed in Table 2.
Landscape descriptors. To calculate the values of
the landscape descriptors, each agricultural eld in the
case study areas was visited, categorised according
to its use and digitised by means of a geographical
information system (GIS).
Four landscape descriptors were calculated in a
200 m radius circle around the observation points
(Table 2).
First, patches (a patch =a relatively homogeneous
non-linear area that differs fromits surroundings) were
assigned to 22 land use types. Three landscape pattern
indices were calculated on the basis of the percentage
cover of the different land use types within the circle:
1. Surrounding habitat variability = number of sur-
rounding land use types (Duelli 1997)
2. Surrounding habitat heterogeneity = number of
patches of different land use types (Duelli 1997)
3. D1 index of landscape pattern (ONeil et al. 1988);
D1 is a measure of dominance:
D1 =ln n + Pi ln Pi, where n is the total number
of land use types and Pi the proportion of patches
in land use type i
Second, a qualitative measurement of landscape
diversity was carried out. The 22 land use types
were aggregated in 4 land use classes to record
information about the landscape quality:
4. Surrounding land use classes: cultivated land, eco-
logical compensation area, forest and built up
area.
Spatial descriptors. Geographical coordinates of the
sites were used as spatial descriptors to detect the
effects of other not measured environmental factors.
Eventual biogeographic or altitude effect (z coordi-
nate) were not supposed to occur at the scale of these
case studies.
Statistical analysis
To collect and retain all the information on the indi-
cators observed, we dened species diversity as both
species variety and relative abundance of the species.
Species-environment relationship was then analysed
with the help of ordinations and multivariate statis-
tics. Multidimensional analysis was rst performed
through the correspondence analysis (CA) by means
of the CANOCO programme (Ter Braak and Smilauer
1998) to obtain ordination diagrams. The result of a
complete linkage clustering was superimposed onto
the CA diagram to separate clusters of objects which
are distinct in dimensions that cannot be represented in
a 2-dimension CA diagram, as proposed by Legendre
and Legendre (1998).
256
Table 1. Strata, habitat types, and the number of sampling sites in the area of region 1
and region 2.
Number of sites
Strata Habitat type Region 1 Region 2
Cultivated areas High intensity meadows 7
Winter wheat elds 20
Extensively meadows 16 3
Ecological Low intensity meadows 7 9
Compensation Hedgerows 3 2
Areas Standard fruit trees in traditional 8
orchards
Wild ower strips 11
Forest edges Forest edges 17 6
To identify the main environmental variables hav-
ing an effect on the species assemblages, a canonical
correspondence analysis (CCA) and a partial CCA,
were carried out (Ter Braak 1996). In CCA, the sig-
nicance of a particular environmental variable can
be assessed by Monte Carlo testing (bootstrapping) of
the axis associated with that variable, using the axis
eigenvalue as the test statistic.
Habitat, landscape and spatial descriptors were in-
troduced in the CCA and partial CCA. Landscape
descriptors were calculated with GIS. To establish a
hierarchy between explanatory variables and to elim-
inate those which do not signicantly explain any
variation, we used CCA with each separate variable
prior to a forward selection, to be followed by CCA
involving all the variables (Jeanneret et al. 1999).
Partitioning of variation was then performed through
partial CCA (e.g., Borcard et al. 1992; Anderson and
Gribble 1998; Pozzi and Borcard 2001). The fraction
of the variation explained (and its signicance, ob-
tained by means of a Monte Carlo permutation test)
by each of the environmental descriptors is given sep-
arately, after eliminating the variation due to the other
(partialed) variables, which are used as covariables.
Results
Faunistic description of the sites
Altogether 16,057 (4 weeks of pitfall-captured) and
15,500 (5 weeks of pitfall-captured) spiders belonging
to 135 and 127 species were collected from the 58 and
51 sites in region 1 and 2, respectively. Altogether,
9,325 (4 weeks) and 32,638 (5 weeks) carabid beetles
belonging to 79 and 96 species were collected from
the 58 and 51 sites in region 1 and 2, respectively.
Due to the unequal sampling effort, the number of
spider and carabid individuals and species of the two
regions should be compared with caution. In both re-
gions, forest edges are well characterised by the spider
and carabid beetle communities and represent a partic-
ular habitat where typical forest species were found
together with species of adjacent meadows (results
published in detail in Jeanneret et al. 2000 and Pffner
et al. 2000).
Altogether, 892 (region 1) and 966 (region 2) but-
teries belonging to 17 and 22 species were observed
on the 58 and 51 sites, respectively. Buttery species
richness was signicantly higher in the extensively
used and low intensity meadows and in the wild ower
strips than in the high intensity meadows and winter
wheat elds (results published in detail in Jeanneret
et al. 2000).
Faunistic comparison of the regions and the habitats
Because of the unequal sampling effort when moni-
toring spiders and carabid beetles in the two regions
(region 1: 4 weeks sampling, region 2: 5 weeks sam-
pling), we tested the time effect (week 1 to week
5) on the species composition of region 2. As the
sampling week missing in region 1 is situated in the
middle of the sampling period (week 4, the last week
of June in 1997) we tested the assumption that one
sampling week situated in the middle of the sampling
period in region 2 would not signicantly change the
species composition. First, we performed CCA with
257
Table 2. Characterisation of the habitats and landscape acting as explanatory variables on
biodiversity.
Scale Environmental variables Land use types
1. Plant species richness
Habitat 2. Habitat type 8 types: High intensity, extensively
descriptors used and low intensity meadow,
hedgerow, standard fruit trees in
traditional orchard
a
, wild ower strip
b
,
winter wheat, forest edge
1. Surrounding habitat variability 22 types: Habitat type + cereal elds,
root crops, corn, rape
b
, vegetable
b
,
pasture, articial meadow, grove, rape,
nursery, slope, brook, built up area,
natural area (forest)
Landscape 2. Surrounding habitat heterogeneity idem
descriptors 3. D1 index of landscape pattern idem
4. Surrounding land use 4 classes: Cultivated land, ecological
compensation area, built up area,
natural area (forest)
Space Coordinate X
descriptors Coordinate Y
a
only in region 1
b
only in region 2
the week as an explanatory variable. On the whole,
the week signicantly affected the species composi-
tion (p 0.005, Monte Carlo permutation test), i.e.,
there is a shift in the species composition and their
relative abundance over the 5 sampling weeks in re-
gion 2. Second, pairwise comparisons were made
and showed that there was no signicant difference
(p =0.94, Monte Carlo permutation test) between the
species composition of weeks 4 and 5. Therefore, the
5 weeks of sampling in region 2 were maintained in
further analysis.
CA ordination diagrams of the sites based on spi-
der, carabid beetle and buttery assemblages differ-
entiate region 1 from 2 (Figures 1, 2 and 3). Never-
theless, superimposition of the results of a complete
linkage clustering shows that spider assemblages of
forest edges and hedgerows of both regions (Figure 1:
cluster 1) show a closer similarity to each other than to
the other habitats of the same region (Figure 1: clus-
ter 2 and 3). Three sites, 1 extensively used, 1 low
intensity and 1 high intensity meadow of region 1 are
exceptions and grouped in cluster 2 with sites of re-
gion 2 (Figure 1). Cluster 3 is exclusively composed of
sites of region 1. For carabid beetles assemblages, like
for spider assemblages, forest edges and hedgerows
of both regions were grouped together, but not the
other habitats of the same region (Figure 2: cluster 1).
Three forest edges and one hedgerow of region 1 were
grouped with meadows of this region (Figure 2: clus-
ter 3). Clusters 2 and 3 are exclusively composed of
sites of region 2 and 1, respectively. For buttery
assemblages, sites of the same region were grouped
together at rst (Figure 3). One cereal eld of region 2
is an exception and belongs to cluster 1 together with
sites of region 1.
Comparison of the habitat, landscape and space
effects on the arthropod groups between the regions
Within the scope of separate CCA and forward selec-
tion procedures, environmental variables and classes
- which explain a signicant part of variation - are
recognised and then introduced in partial CCA. The
habitat descriptors act differently according to arthro-
pod groups and regions (Table 3). Plant species
richness explains a signicant part of the variation
for every arthropod assemblage in region 1 (spiders:
2.5%; carabid beetles: 2.6%; butteries: 3.8%). In
both regions the habitat type is a signicant explana-
tory variable for epigeal arthropod assemblages. For
butteries in region 2, signicance is only achieved by
258
Figure 1. CA ordination of the 58 sites (region 1) and 51 sites
(region 2) based on spider assemblages. The ellipses represent the
result of a complete linkage clustering. r1, r2 = sites belonging to
the region 1 and region 2, respectively. For visual clarity, member-
ship of well gathered sites is indicated by Region 1 or Region 2.
The arrow indicates that the site belongs to region 2 and cluster 2.
adding the variation explained by plant species rich-
ness and habitat type. On the one hand, landscape
descriptors have no inuence on the arthropod as-
semblages, if calculated as an index summarising the
information like surrounding habitat variability and
heterogeneity, and D1. On the other hand, if the land
use classes are taken into account, spider assemblages
of region 2 are signicantly inuenced by the presence
of both natural areas and ECA in the surroundings of
the habitat where they were caught, carabid beetles
react signicantly to both cultivated land and natural
area in region 1, and buttery assemblages are sig-
nicantly inuenced by both natural area and ECA
in region 1. Spatial position of the sites is a signif-
icant explanatory variable for the epigeal arthropods
in both regions, but only in region 1 for the buttery
assemblages. The habitat type is the most inuencing
factor for epigeal arthropods in both regions, while
surrounding land use is more important for butteries
in region 1. In region 2, when tested alone, no environ-
mental variable measured explains any part of buttery
assemblages.
Figure 2. CA ordination of the 58 sites (region 1) and 51 sites (re-
gion 2) based on carabid beetles assemblages. The ellipses represent
the result of a complete linkage clustering. r1, r2 = sites belonging
to the region 1 and region 2, respectively. For visual clarity, member-
ship of well gathered sites is indicated by Region 1 or Region 2
The arrows indicate that the sites are grouped with the meadow sites
of region 1.
The large amount of unexplained variation is due
to factors overlooked in this study or to stochastic
variation.
Discussion
Methodological aspects
To estimate the total number of spider and carabid
beetle species occurring in our regions, pitfall traps
should be operated for a longer sampling period than
in this study. Nevertheless, Duelli (1990) showed that
in comparison with a full season sample of 28 weeks,
more than 70% of the number of species is obtained
in similar habitats in Switzerland within 4 sampling
weeks from the beginning of May to the beginning of
July for both spiders and carabids. Depending on the
habitat, the percentage of species caught ranged from
259
Table 3. Summary of the percentage variation explained by environmental variables for spiders, carabid beetles, and
butteries in two regions after partitioning with partial CCA. 1 = region 1 and 2 = region 2. When given, percent of
variation is signicant at p < 0.05 (Monte Carlo permutation test). n.s.: not signicant.
Spiders Carabid Butteries
beetles
Region 1 2 1 2 1 2
Scale Environmental variables Explained variation (%)
Habitat
descriptors
Plant species richness
Habitat type
2.5
16.6
n.s.
14.2
2.6
9.8
n.s.
21.9
3.8
n.s.
11.1
Surrounding habitat variability n.s. n.s. n.s. n.s. n.s. n.s.
Surrounding habitat heterogeneity n.s. n.s. n.s. n.s. n.s. n.s.
D1 index of landscape pattern n.s. n.s. n.s. n.s. n.s. n.s.
Landscape Surrounding land use classes:
descriptors
Cultivated land
Natural area (forest)
Ecological compensation area
n.s.
n.s.
n.s.
n.s.
4.2
4.4
n.s.
n.s.
n.s.
n.s.
n.s.
6.8
n.s.
n.s.
n.s.
Built up area n.s. n.s. n.s. n.s. n.s. n.s.
Space 4.9 4.0 4.4 3.5 6.4 n.s.
Total of the variation explained
a
26.3 28.6 25.3 33.2 17.4 11.1
a
The total takes into account the fraction of the variation explained in common between the environmental variables.
84.6% (winter wheat) to 86.0% (meadows) for cara-
bids and from 73.5% (winter wheat) to 77.8% (mead-
ows) for spiders (Duelli 1990). Within the framework
of our study, these differences are acceptable and al-
low the comparison between the habitat types. Duelli
(1990) also showed that an additional sampling week
during the optimum period (MayJuly) results in
less than 10% of the species. As mentioned in Sec-
tion Regions, sampling methods, sampling in cereal
elds for a longer period of time would cause dras-
tic habitat changes by harvesting and sowing of the
next crop. These changes would strongly affect the
comparison between habitat types. Nevertheless, for
carabid beetles a short sampling period in May-June
causes a bias when comparing forest with open habi-
tats because species active in spring are more frequent
in open habitats than in forests which are characterised
by species active in autumn (anonymous reviewer,
personal communication). However, hedgerows and
forest edges are a part of the open cultivated land-
scape in our regions. The comparison with pure
forests would be more problematic. Furthermore, in
our study spider and carabid beetle assemblages dif-
fered considerably in the open habitats and the forest
edges as well as hedgerows (CA ordinations) despite
the probable underestimation of the species richness in
the latter. Sampling all over the year would show the
differentiation of both types of habitats more acutely.
Examining species assemblages allows a compre-
hensive appreciation of the impact of habitat and
landscape on biodiversity. We have used this approach
instead of summarising the biotic information in one
single value such as species richness or a diversity
index where interpretation would be difcult and the
loss of information too substantial.
As emphasised by Anderson and Gribble (1998),
we cannot suggest that the methodology of variance
partitioning would allow the establishment of any
causal effects, which would require proper experimen-
tal design and analyses. The method corresponds to
the univariate multiple regression, including its warn-
ings concerning issues of causal relationships, choice
of variables and redundancy of parameters causing in-
creases in explained variation due to chance alone.
Nevertheless, the landscape and its components should
be included in the environmental control model as ex-
planatory variables, as it has been demonstrated in this
study. Thus, the search and analysis of correlations
is certainly the best methodology to be used because
deliberate experiments which would result in manip-
ulating the landscape in an experimental design are
practically unfeasible.
260
Figure 3. CA ordination of the 58 sites (region 1) and 51 sites (re-
gion 2) based on buttery assemblages. The ellipses represent the
result of a complete linkage clustering. For visual clarity, member-
ship of well gathered sites is indicated by Region 1 or Region 2.
The arrow indicates that the site is grouped with sites of region 1.
Landscape and habitat features as environmental
control factors
CA ordination diagrams associated with clustering re-
sults show that regions 1 and 2 have their own specic
pool of spider, carabid beetle and buttery species,
but the difference between the forest edges and the
other habitats is greater than the difference between
the regions for epigeal arthropods.
The correlative models obtained from partial CCA
allow to discriminate between both components of the
environmental control, i.e., habitat and landscape, and
space. On the one hand, spider and carabid beetle as-
semblages are inuenced by the habitat type. This is
consistent with other studies on the characterisation
of habitats with spiders (e.g., Duffey 1974; Clausen
1986; Alderweireldt 1989; Martin 1991) and cara-
bid beetles (e.g., Luff et al. 1989; Turin et al. 1991;
Kramer 1996). In both regions, forest edges are char-
acterised by specic assemblages. The relationship is
particularly direct between the carabid beetle assem-
blage and the habitat type in region 2. In this region
of arable elds, the habitat type gradient, in other
words the difference between the habitats, is greater
than in the meadow landscape (region 1), and this
plays an important role for less mobile organisms like
carabid beetles in comparison with spiders since less
exchanges with neighbour elds occur. Contrary to
that, spiders assemblages characterise the habitat type
irrespective of the habitat type gradient.
On the other hand, it is surprising that the habi-
tat type plays such a minor role for butteries since
it signicantly explains the species composition only
in conjunction with the oristic richness in region 2,
although the obviousness of the relationship has been
demonstrated in the literature (e.g., Dennis 1992; Kre-
men 1992; Debinski and Brussard 1994). Our explana-
tion is that the very poor buttery species assemblages,
17 and 22 species in region 1 and 2, respectively,
are mainly composed of generalist species as dened
by Ouin and Burel (2002), considering their disper-
sal ability (Warren 1992) and degree of polyphagy
(Scriber 1973). Generalists species are less infeoded
to a particular habitat and therefore, the habitat type
has less inuence.
Plant species richness is a signicant explanatory
factor for epigeal arthropods in region 1 and in both
regions for butteries. For epigeal arthropods, this
may be explained by the species pools which are dif-
ferent in both regions. The spider and carabid beetle
assemblages captured in region 1, which is domi-
nated by meadow ecosystems, are sensible to the
habitat structure, which is represented by the plant
species richness, while assemblages captured in re-
gion 2, which is dominated by arable elds, are not
signicantly inuenced by this factor.
As nectar feeding insects, butteries strongly
depend on owering plants. Distribution of nectar
sources in space and time plays a crucial role in the
location and dispersal of buttery populations (Boggs
1987). Habitats with higher plant species richness of-
fer a nectar source which is better distributed in time
than habitats with fewer species. Therefore, habitats
rich in plant species are occupied and visited by a
larger buttery species set.
As stressed by Wagner and Edwards (2001), habi-
tat variability and heterogeneity are simple to quan-
tify, but depend on the habitat classication and it
261
is assumed that all habitat types are equally differ-
ent from each other so that the specic composition
of a landscape does not matter. Our study shows that
the specic composition of the landscape does mat-
ter and that the loss of information resulting from
simple index calculations is too substantial and as a
consequence no signicant part of the variation in
spider, carabid beetle and buttery assemblages can
be explained. If its specic composition is taken into
account, the surrounding landscape becomes a signif-
icant explanatory factor for spiders and butteries of
region 2 (natural area + ECA), and carabid beetles of
region 1 (cultivated land + natural area).
In the region where the species assemblages are
less inuenced by the habitat type, the landscape
composition is a stronger explanatory factor, i.e., re-
gion 2 for spiders and region 1 for carabid beetles and
butteries.
For spiders, these results do not conrm previ-
ous studies carried out by Asselin and Baudry (1989),
Burel and Baudry (1995) showing no effect of the
landscape structure. Our results show that particular
habitats like ECA and natural areas in the surround-
ings may control the attainability of the habitat for
spiders.
In our study landscape descriptors and the sur-
rounding habitat type in particular have no major
inuence on buttery assemblages in one of the re-
gions as postulated for some groups by Dover et al.
(1992) and as seen from results for particular species
(Thomas and Harrison 1992; Thomas and Hanski
1997). Most buttery species y over the landscape,
visiting small or large areas. They need structure to
move and often require several habitats to complete
their life-cycles. Therefore, butteries should be in-
uenced by the habitat arrangement around a point
that they visit. In region 2, however, the lack of ver-
tical structures like hedgerows, forest edges, ditches,
etc. leads to a uniform attainability of the habitats
for butteries. Once again, an analysis of the assem-
blages focussed on functional groups should reveal
differences between generalist and specialist species.
The range of variation explained by spatial vari-
ables indicates that other factors that were not tested
could play a role. In particular, the action of a mi-
croclimatic shift is possible in area 1 where slopes
exposed to the north and therefore wetter and colder,
are mixed with drier and warmer places exposed to the
south. Furthermore, other landscape features like the
fractal dimension, the contagion index (ONeil et al.
1988) and structural features dened by Marino and
Landis (1996) as well as connectivity measurements
should be added in the model. An additional explana-
tory factor which was not included and which could be
signicant is the history of the sites. Indeed, while at
the same time the habitats are currently managed in the
same manner, their history can have an importance in
the determination of the assemblages of species which
we observe in the present.
Conclusions
Because of the differentiated response shown by the
arthropod taxa to habitat and landscape features, it is
important to approach the environmental control by
examining different taxa particularly when the goal is
to evaluate restoration programs or to found manage-
ment recommendations in agricultural landscapes.
As it has been demonstrated in this study, land-
scape and its components should be included in
the environmental control model as explanatory vari-
ables. Nevertheless, the complex relationship between
arthropods and landscape in a multi-indicator ap-
proach is fragmentary in spite of studies in recent
years (e.g., Burel and Baudry 1995; Paoletti 1999;
McCracken et al. 2000; Atauri and de Lucio 2001;
Tscharntke et al. 2002; Weibull 2002) and in com-
parison with the links to habitat features. Especially,
temporal uctuations of arthropod abundance are well
known and inuence the analysis of the impact of en-
vironmental factors. Medium- and long-term studies
are necessary to analyse the temporal trends and to
separate them from the other effects. Our study will
contain 4 sampling years between 1997 and 2004 and
will therefore allow comparisons in the medium term.
Furthermore, as the response of the taxa to the habi-
tat and landscape features depends on the region, data
should be collected in other regions across Europe to
maximize landscape gradient. Then results could be
compared to allow generalization.
Acknowledgements
This study was partly nanced by the Swiss Federal
Ofce for Agriculture. The authors wish to thank J.
Steiger, G. Blandenier and H. Hnggi for spider iden-
tication, H. Luka for carabid beetle identication, S.
Bosshart, M. Waldburger for their assistance in eld
work.
262
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