Agriculture, Ecosystems and Environment 78 (2000) 215–222

Overwintering of arthropods in soils of arable fields and

adjacent semi-natural habitats
Lukas Pfiffner ∗ , Henryk Luka
Research Institute of Organic Agriculture (FiBL), Ackerstrasse, CH-5070 Frick, Switzerland
Received 11 March 1999; received in revised form 12 August 1999; accepted 22 September 1999

Abstract
In order to determine the significance of field margins for the overwintering of arthropods in agricultural landscapes,
different sites of an integrated and of an organically managed farm were investigated in the northwest of Switzerland. During
December 1995 and January 1996, soil samples were taken with an electronic-hydraulic soil borer (depth: 25 cm, diameter:
8 cm). After hand sorting the larger arthropods, the small ones were extracted with a modified MacFadyen apparatus.
The abundance of arthropods in the arable fields was significantly lower than in the adjacent semi-natural habitats. Highest
abundances and species diversities were found in a sown wildflower strip, a hedge, a permanent meadow and a meadow
under the cherry trees of the organic farm. With a total of 90 arthropod species in the semi-natural habitats, five times more
species were found than in the arable fields. Staphylinids, carabids, spiders and chilopods were the most abundant arthropod
groups. The data showed that undisturbed semi-natural habitats and extensively managed field margins play a key role as
overwintering sites for many predatory arthropods. ©2000 Elsevier Science B.V. All rights reserved.
Keywords: Overwintering; Arable land; Field margins; Semi-natural habitats; Beneficial arthropods; Switzerland

1. Introduction of intensive farming methods have been documented

In agricultural landscapes there is a general trend
towards a reduction in diversity and abundance of
native flora and fauna because of increased landscape
uniformity and greater disturbance via farm inputs,
cultivation and management (Burel, 1992; Pfiffner,
1997). Large disturbances, by both physical and
chemical means, can reduce abundance and diversity
of beneficial species, and ecosystem functions can be
altered (Giller et al., 1997). Declines in the distribu-
tion and abundance of beneficial arthropods because
∗ Corresponding author. Tel.: +41-62-865-72-72;
fax: +41-62-865-72-73.
E-mail address: lukas.pfiffner@fibl.ch (L. Pfiffner).
in cornfields (Dritschilo and Erwin, 1982), potatoes
(Kromp, 1989) and wheat (Pfiffner and Niggli, 1996).
Since 1993, the Swiss government has subsi-
dized sustainable farming methods including organic
farming and the establishment of field margins and
semi-natural habitats. One of the objectives of this
subsidy is to halt the marked decline of biological
diversity in agricultural landscapes. It is thus im-
portant to evaluate the effects of diversity-enhancing
techniques on cultivated land and its surroundings.
Field margins are important refuges for many
plants and animals and thus play a key role in main-
taining biological diversity on farmland (Fry, 1994).
The size and type of field margins and associated
semi-natural habitats play an essential role in the

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216 L. Pfiffner, H. Luka / Agriculture, Ecosystems and Environment 78 (2000) 215–222

survival of many invertebrates during winter. This has
been shown for polyphagous pest predators, such as
carabids, staphylinids and spiders (Sotherton, 1984;
Wiedemeier and Duelli, 1993; Andersen, 1997).
In arable land, these beneficial epigeic arthropods
are important to reduce various pests (Cocquempot
and Chambon, 1990; Symondson, 1993; Menalled
et al., 1999). However, there is a lack of informa-
tion about appropriate overwintering sites and the
effects of different field margins on several groups of
invertebrates.
The objective of this study was to compare the
overwintering arthropod populations in arable fields
with those in different adjacent semi-natural habitats
and field margins (comparison in pairs). Two different
landscapes were examined in the northwest of Switzer-
land: a recently improved, integrated managed arable
landscape with grassy field margins and since 1974 an
organically managed farm in a richly structured land-
scape.
2. Materials and methods
2.1. Study site
The climate at the study sites is relatively mild and
dry with a mean precipitation of 785 mm per year and a
mean temperature of 9.5◦ C. The altitude is 350–365 m
above sea level. The soil on both farms is a Luvisol
from loess with similar soil properties.
On the integrated managed farm at Aesch, two win-
ter wheat fields and two adjacent meadow strips were
chosen (Table 1): The meadow strips 5 m in width (un-
fertilized and mown twice per year) as well as hedges
along the arable fields, were established 5 years ago to
enrich the impoverished landscape. These strips were
sown with a grass-clover mixture with 14 additional
wildflowers (e.g., Centaurea jacea, Silene vulgaris,
Chrysanthemum leucanthemum).
On the organic farm at Oberwil, in a more diver-
sified landscape, the following habitats were exam-
ined (Table 1): a winter wheat, an arable meadow,
a ploughed wildflower strip as control plot (tilled
strip without vegetation), and adjacent semi-natural
habitats (a 2-year-old wildflower strip, a hedge, stan-
dard cherry trees on a permanent ‘Arrhenatheretum’
meadow). The arable meadow was a temporary ley
of legumes (Trifolium pratense, T. resupinatum, T.
alexandrinum and Medicago sativa) and grass (Lolium
multiflorum). The highest plant diversity was found
in the 3 m wide wildflower where a plant mixture of
25 species was sown (e.g., endangered species such
as Agrostemma githago, Centaurea cyanus, Legousia
speculum-veneris, Echium vulgare; for a full list see
Nentwig, 1995). This strip had lain fallow for the
two previous years. For further characteristics of the
investigation sites see Table 1.

Table 1
Main characteristics of the investigation sites
Arable plots in the rotation Abbreviation Agea Farming Previous crop
Winterwheat WW-1 1 integrated sunflowers
Winterwheat WW-2 1 integrated rape
Winterwheat WW-3 1 organically potatoes
Arable meadow (grass-legumes mixture) AM 1 organically barley
‘Control’ plot for the wild flower strip Con 0.5 plouged strip in autumn wildflower strip

Adjacent field margins and semi-natural habitats Abbr. Agea Plant div.b Width (m) Management

Sown meadow strip (grass-weed-flower mixture) SM-1 5 15 4–5 2–3 cuttings per year
Sown meadow strip (grass-weed-flower mixture) SM-2 5 26 4–5 2–3 cuttings per year
Permanent meadow strip (Arrhenateretum) PM >15 23 4–5 2 cuttings per year
Cherry trees on a permanent meadow CT >15 24 6–7
hedge H >16 16 3–4 1 cutting per 5 years
Sown wildflower strip (25 species sown) WS 2 44 3 no cultivation since 2 years
a Years.
b Number of plant species.
 L. Pfiffner, H. Luka / Agriculture, Ecosystems and Environment 78 (2000) 215–222 217

2.2. Sampling of arthropods
On three dates between December 1995 and January
1996, soil samples were taken at six paired habitats
(arable or control plot and adjacent field margin) on the
two farms described above. Twenty-four undisturbed
soil cores were taken with an electronic-hydraulic soil
borer (depth: 25 cm, diameter: 8 cm), randomly leav-
ing about 8 m space between two consecutive samples
at each site. Samples in the arable plots were taken
in a line parallel to and 30 m distant from the field
margins. After manually sorting the larger arthropods,
the samples were placed in a modified MacFadyen
apparatus for 4 days to extract the small arthropods
(Bieri et al., 1978). Most arthropods were determined
to species level including carabids larvae and juvenile
spiders so far as possible. Finally, the 24 samples from
each habitat were pooled.
2.3. Statistical methods
In each case the data showed that the arthropods
were highly aggregated. Therefore, statistical analy-
ses were done with pooled data from the three sam-
pling dates. The comparison of paired arable field and
adjacent semi-natural habitat was conducted with the
Wilcoxon rank test. Arthropod community classifica-
tion was established using an agglomerative hierar-
chical clustering with the minimum variance method
of Ward (Orloci, 1967; JMP Version 3). The species
richness of each habitat was compared using an ordi-
nation of the JACCARD similarity index with multi-
dimensional scaling (NTSYS 1.8, Rohlf, 1993).
habitats at each site. Highest abundances and species
diversities were found in the sown wildflower strip,
hedge, permanent meadow and under the cherry trees
on the organic farm (Fig. 1).
The more frequently found arthropod groups were
pest predators, such as staphylinids, carabids, spiders
and chilopods. Furthermore, large numbers of aggre-
gated staphylinids were observed by a visual control
near the south exposed parts of the cherry tree stems.
Ninety of the 94 arthropod species found in all habitats
were in semi-natural habitats; there were 18 species
in the five arable fields; 75 species were exclusively
found in the semi-natural habitats; and only four ex-
clusively in the arable fields (Table 2).
Fig. 2 shows the community composition of the
arthropods: the most even composition of the arthro-
pod groups was in the sown meadows, hedge and
wildflower strip. The staphylinids were the most
abundant group in nearly all habitats. This domi-
nance was particularly marked in the arable plots.
The diplopods might be enhanced in wheat Plot 1
and in the 2-year-old wildflower strip because of rich
litter layers there. In wheat Plot 1 this was because of
a previous sunflower crop, which left much organic
material on the soil.
Cluster analysis showed that the arthropod popula-
tions of the semi-natural habitats on the organic farm
were similar. All arable field samples formed one clus-
ter, and the two sown grass-weed margins were in be-
tween (Fig. 3). The higher species richness of field
margins and their relative similarities were expressed
in the two main clusters using an ordination of the
JACCARD similarity index (Fig. 4).

Table 2
3. Results Total number of found arthropod species in arable land and adjacent
field margins. Numbers in brackets are numbers of exclusively
found species in the corresponding landscape category
The abundance of arthropods varied from 33 to 1163
Arthropods Arable landa Field margins Total
individuals per m2 (Fig. 1). Lowest abundances and (n = 5) (n = 6)
species diversities were generally found in the arable
Carabidae 4 (1) 22 (19) 23
fields (33–107 ind./m2 , 3–6 species). On the other Staphylinidae 7 (2) 36 (30) 38
hand there was a wide range in the semi-natural habi- Coccinellidae 1 (1) 1 (1) 2
tats (134–1163 ind./m2 , 13–37 species). In all cases Araneae 1 (0) 17 (16) 17
the abundance of arthropods in the arable fields was Chilopoda 2 (0) 5 (3) 5
significantly lower than in the adjacent semi-natural Diplopoda 3 (0) 6 (3) 6
Isopoda 0 (0) 3 (3) 3
habitat, with only one exception (Wilcoxon rank test,
Fig. 1). No differences in abundance were found Total 18 90 94
a
within the arable fields and within the semi-natural Including control plot.
218 L. Pfiffner, H. Luka / Agriculture, Ecosystems and Environment 78 (2000) 215–222

Fig. 1. Overwintering arthropods in arable fields and adjacent semi-natural habitats on two farms. (a) Densities of arthropod per m2 ;
(b) number of arthropod species. Wilcoxon rank test, *: p < 0.05, **: p < 0.01, ***: p < 0.001. Abbr. see Table 1.

4. Discussion fields, more perennial crops and more semi-natural

The results show that field margins which are not
ploughed or otherwise disturbed during autumn or
winter can act as refuges and hibernation sites for
many arthropods. In the less intensively cultivated and
more structured landscape of the organic farm, the
abundance of arthropods was two to five times greater
than in the plots of the integrated farm. The landscape
of the organic farm was more diversified: smaller
habitats lead to a higher spatial complexity. Nonethe-
less, plant diversity and the management of the mead-
ows were very similar at most sites of both farms. The
higher landscape complexity and age of the mead-
ows on the organic farm might be the main reason
for their better performance (in terms of abundance
and species richness) compared with the meadows
of the integrated farm. Higher agroecosystem comp-
lexity and low input agriculture (organic farming)
 L. Pfiffner, H. Luka / Agriculture, Ecosystems and Environment 78 (2000) 215–222 219

Fig. 2. The community composition of the main arthropod taxa in arable fields and the adjacent semi-natural habitats. Abbr. see Table 1.

can positively influence the fecundity and feeding of
beneficial arthropods (Bommarco, 1998). Bommarco
found significant larger ground beetles (Poecilus
cupreus) and a three times higher fecundity in a more
complex and organically managed landscape than in
a monotonous, conventionally farmed landscape in
Sweden.
The mean density of more than 1100 arthropods per
m2 found in the extensive meadow under the cherry
trees is relatively high. Sotherton (1985) considered
densities of 1000 epigeic polyphagous predators in
field boundaries to be very high.
The 2-year-old, sown wildflower strips which were
undisturbed (uncultivated) for 2 years contained the
highest number of arthropod species. This type of
semi-natural habitat is highly attractive to many arthro-
pod species of arable land because of a high botan-
ical and structural diversity, the presence of annual,
biennual and perennial plant species and a rich
litter layer. The non-cultivation and the favourable

Fig. 3. Arthropod community classification using an agglomerative hierarchical clustering with the minimum variance method of Ward.
Abbr. see Table 1.
220 L. Pfiffner, H. Luka / Agriculture, Ecosystems and Environment 78 (2000) 215–222
Fig. 4. The species richness of each habitat using an ordination of the JACCARD similarity index with multidimensional scaling. Abbr.
see Table 1.
botanical characteristics of this strip would appear to
be among the key factors. In addition, the colonization
of diverse field margins by arthropods can be fairly
rapid, as shown in the investigation by Thomas et al.
(1994). In a similar study, Lys and Nentwig (1994)
found a significantly higher density and species rich-
ness of overwintering carabids, staphylinids and spi-
ders (mean density of 1300 ind./m2 ) in a 2-year-old
wildflower strip in comparison with an adjacent crop
of winter barley.
High densities of overwintering polyphagous preda-
tors have often been found in dense vegetation (e.g.,
Sotherton, 1985; Thomas et al., 1991). Furthermore,
Desender (1982) showed a positive relation between
the mean depth of sod layer in grass and the den-
sities of overwintering carabids. The presented data
from the ploughed wildflower strip (control plot with-
out vegetation) showed that the vegetation was of de-
cisive importance for the hibernation site selection
of many arthropods. One of the most important fac-
tors inducing high densities of polyphagous predators
seems to be the density or biomass of the vegetation
layer; Bürki and Hausamann (1993) observed lowest
densities of arthropods under plant species with lit-
tle vegetation cover such as Agrostemma githago or
Chenopodium polyspermum and very high densities of
predators (more than 2100/m2 ) under highly attractive,
larger plants such as Achillea millefolium and Arctium
minus. These two last mentioned plant species were
only present in the wild flower strip.
The permanent, undisturbed vegetation of semi-
natural habitats can reduce diurnal temperature fluctu-
ations and diminish the mortality rates of arthropods
in the uppermost soil of wild flower margins, as
Bürki and Hausamann (1993) have shown. During
the present sampling period the soil cores of the
arable fields were mostly frozen down to 5 cm deep,
which was never observed in adjacent semi-natural
 L. Pfiffner, H. Luka / Agriculture, Ecosystems and Environment 78 (2000) 215–222
habitats. The temperature buffering properties of the
semi-natural habitats could be a key abiotic factor in
overwintering site selection.
Field margins can act as reservoirs for polyphagous
arthropods. Indeed, during the overwintering pe-
riod, top epigeic predators such as some staphylinid
species (Paederus fuscipes, P. litoralis, Tachyporus
chysomelinus, T. hypnorum, Xantholinus linearis, X.
longiventris), ground beetles (Bembidion lampros,
Agonum muelleri, Demetrias atricapillus) and 16
spider species were only found in the field margins.
During the vegetation period, most of these species
were also found in the adjacent arable fields (data
not published). This contrasts partly with the findings
of Andersen (1997) in Norway. On the one hand,
he showed that most carabid and staphylinid species
were significantly more abundant in field margins
than in the adjacent cereal fields. On the other hand,
he concluded that the grass field margins were of lim-
ited importance as a reservoir for predatory beetles,
because he found the ground beetle Clivina fossor
and staphylinids of the genus Lathrobium distributed
evenly over field margins and cereal fields.
In addition, Hawthorne et al. (1998) showed that un-
cropped wildlife strips could enhance the abundance
of polyphagous predators within the field because they
can act as an additional source of colonizing individu-
als (e.g., Bembidion lampros) and thus do not create a
barrier to the movement. This fact underlines their rel-
evance for the biological control of pests in adjacent
fields. However, there are data of Mauremootoo et al.
(1995) which show reducing effects of hedgerows on
the dispersal of carabid beetles between fields.
Another relevant feature is the spatial distribution
of arthropods within field margins. Riedel (1995)
found a preference by field inhabiting carabids such as
Bembidion lampros and B. tetracolum as well as for all
pooled carabids for the edge (ecotone) of uncultivated
grass vegetation. No significant distribution preference
was observed for Amara familiaris, A. plebeja and
Tachyporus hypnorum. These flying and larger species
are known to overwinter often in the hedgerows.
5. Conclusions
Semi-natural habitats are important overwintering
sites for arthropods and can significantly enhance the
221
number of species and the abundance of beneficial
arthropods on arable land. They can fulfill multiple ob-
jectives in agroecology and nature conservation. This
is encouraged by structural complexity, botanical di-
versity and the permanent vegetation layer in these
habitats.
Acknowledgements
The authors thank Dr. A. Haenggi (Natural History
Museum, Basel) for use of the MacFadyen apparatus;
T. Blick (spiders), Dr. C. Huber (carabids larvae),
J. Spelda (myriapods), S. Kiener (staphylinids) for
species determinations and the farmers for providing
their fields; and the Basellandschaft canton adminis-
tration and the Swiss Agency for the Environment,
Forests and Landscape for financial support. Thanks
also go to Prof. Dr. P. Duelli, Dr. B. Speiser, Dr. P.
Mäder and P. Damary for their helpful comments on
the manuscript and the anonymous referees and an
Editor-in-Chief for improvements to an earlier version
of this paper.
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