ARTICLE IN PRESS

Basic and Applied Ecology 9 (2008) 682–690 www.elsevier.de/baae

Insect pests in winter oilseed rape affected by field and

landscape characteristics
Johann G. Zaller!, Dietmar Moser, Thomas Drapela, Claudia Schmöger, Thomas Frank
Department of Integrative Biology and Biodiversity Research, Institute of Zoology, University of Natural Resources and Applied
Life Sciences, Gregor Mendel Strasse 33, A-1180 Vienna, Austria

Received 24 January 2007; accepted 18 October 2007

Abstract
Winter oilseed rape (OSR, Brassica napus) cropping is often associated with an intensive use of pesticides. The
transformation of structurally rich landscapes into more monotonous landscapes may be partly responsible for this,
because non-crop habitats believed to benefit natural enemies have been eliminated. We examined the influence of field
(soil quality, nitrogen fertilization) and landscape characteristics (OSR area and isolation, non-crop area, landscape
diversity, proportions of grassy fallows and woody areas) on three major European OSR pest groups: pollen beetles,
stem weevils, and brassica pod midges. Twenty-nine landscape sectors ranging from structurally poor to complex were
studied at eight spatial scales (radii 250–2000 m) centered in the studied OSR fields. Abundances of pollen beetles and
stem weevils were significantly positively correlated with soil quality and negatively related to OSR area in the
surroundings. Generally, abundances of all groups were positively related to woody areas, but not related to grassy
fallow area. Pod midges and stem weevils tended to respond primarily to landscape variables at small (250–500 m) and
medium (1000–1250 m) scales, while pollen beetles responded at medium to large (1000–2000 m) scales. The results are
discussed in relation to differences in overwintering strategies and mobility of pest insects. Strategies at the field and
landscape level, aiming to reduce pest pressure in OSR fields, are also discussed.
r 2007 Gesellschaft für Ökologie. Published by Elsevier GmbH. All rights reserved.

Zusammenfassung
Der Anbau von Winterraps (OSR, Brassica napus) ist oft mit einem intensiven Pestizideinsatz verbunden. Zumindest
teilweise für diese Entwicklung verantwortlich gemacht wird die Umgestaltung von strukturreichen zu monotonen
Landschaften, da dabei naturnahe Habitate verloren gehen, die für die natürlichen Gegenspieler der Schädlinge
förderlich sind. Wir untersuchten den Einfluss von Feld- (Bodenqualität, Stickstoffdüngung) und Landschaftscha-
rakteristika (Anteil und Isolation der Rapsflächen, naturnahe Flächen, Landschaftsdiversität, Anteil an Grasbrachen
und Gehölzflächen) auf die Abundanzen von drei in Europa wichtigen Gruppen von Rapsschädlingen:
Rapsglanzkäfer, Gefleckter Kohltriebrüssler und Großer Rapsstängelrüssler und Kohlschotenmücke. An 29, von
strukturarm bis komplex reichenden Landschaftssektoren wurden auf acht konzentrischen Skalen um das
Untersuchungsfeld (Radius 250–2000 m), die Beziehungen zwischen Schädlingsabundanzen und Feld-/Landschaftvar-
iablen untersucht. Die Abundanzen der Rapsglanzkäfer und der Stängelrüssler korrelierten signifikant positiv mit der
Bodenqualität und signifikant negativ mit der Rapsfläche in der Umgebung. Generell waren die Abundanzen aller
!Corresponding author. Tel.: +43 1 47654 3205; fax: +43 1 47654 3203.
E-mail address: johann.zaller@boku.ac.at (J.G. Zaller).

1439-1791/$ - see front matter r 2007 Gesellschaft für Ökologie. Published by Elsevier GmbH. All rights reserved.
doi:10.1016/j.baae.2007.10.004
 ARTICLE IN PRESS
J.G. Zaller et al. / Basic and Applied Ecology 9 (2008) 682–690 683

Schädlingsarten positiv beeinflusst von der umgebenden Waldfläche aber unbeinflusst vom Anteil an grasigen Brachen.
Kohlschotenmücken und Stängelrüssler reagierten vorwiegend auf Landschaftsvariablen im näheren (250–500 m) und
mittleren Umfeld (1000–1250 m), während Rapsglanzkäfer auf mittleren bis großen Skalen reagierten (1000–2000 m).
Die Resultate werden in Bezug zu den unterschiedlichen Überwinterungsstrategien und der Mobilität der Schädlinge
diskutiert.
r 2007 Gesellschaft für Ökologie. Published by Elsevier GmbH. All rights reserved.

Keywords: Canola; Ceutorrhynchus napi; Ceutorrhynchus pallidactylus; Dasineura brassicae; Insect pests; Landscape ecology;
Meligethes aeneus; Oilseed crop; Pest–crop interactions; Spatial scales

Introduction Pest species considered in the current study include

In agroecological research it has been appreciated that
plant–insect interactions and other ecological processes
depend on scales much larger than a single habitat
(Levin, 1992; Schneider, 1994; Wiens, 1989). Moreover,
studies using a landscape approach where area propor-
tions, spatial arrangement and connectivity of habitats
were considered have shown that the influence of
landscape complexity and structure tends to be spe-
cies-specific and communities are made up of species
with different spatial strategies (Kareiva & Wennergren,
1995; Pickett & Cadenasso, 1995; Thies & Tscharntke,
1999; Wiens, Schooley, & Weeks, 1997; With, Pavuk,
Worchuck, Rhonda, & Fisher, 2002). Crop–pest inter-
actions have mainly been studied on single-pest species
by focusing either on the impact of field parameters or
on landscape structure but only rarely included both
factors (Östman, Ekbom, & Bengtsson, 2001a). We
investigated how the abundances of three major func-
tional groups of insect pests in winter oilseed rape (OSR,
Brassica napus L., canola) responded to field parameters
and landscape characteristics at various spatial scales.
We are not aware of any other studies that have
examined insect pest–field/landscape interactions on
more than one group of insect pests.
The importance of OSR as a source for industrial and
nutritional oil has been increasing worldwide during the
last decades as reflected by an increase of its acreage
from 8.2 million ha to 27.0 million ha from 1970 till 2005
(http://faostat.fao.org/site/567/default.aspx). In many
regions this is accompanied by a dramatic dispropor-
tionate increase of pesticide application (Gianessi &
Marcelli, 2000). Prevalent pest problems are often
attributed to the transformations of formerly hetero-
geneous landscapes with high proportions of semi-
natural non-crop habitats to more monotonous landscapes
mainly dominated by arable land and its local decrease
of biodiversity (Kareiva, 1990; Pickett & Cadenasso,
1995) and detrimental consequences for interactions
between pests and beneficial organisms (Menalled,
Marino, Gage, & Landis, 1999; Östman, Ekbom,
Bengtsson, & Weibull, 2001b; Polis, Anderson, & Holt,
1997; Roland & Taylor, 1997; Thies & Tscharntke,
1999).
(i) stem weevils (Ceutorhynchus pallidactylus Marsh. and
Ceutorhynchus napi Gyll., Coleoptera, Curculionidae) that
lay eggs in leaf petioles or midribs of OSR plants while the
larvae tunnel in the stems; (ii) pollen beetles (Meligethes
aeneus Fabr. and M. viridescens Fabr., Coleoptera,
Nitidulidae) that feed on pollen and destroy flower buds
and (iii) brassica pod midge (Dasineura brassicae Winn,
Diptera, Cecidomyiidae) that lay eggs into OSR pods
where the hatched larvae consume the seeds as well as
tissue of the pod walls and cause the pods to split
prematurely (Alford, Nilsson, & Ulber, 2003). Stem
weevils either pupate in the soil of the OSR field or in
woodland and grassy field boundaries and are not very
mobile; the pod migdes have more than one generation a
year and pupate in the OSR field from which the weakly
flying adults invade new OSR fields of the next year; the
very mobile pollen beetles hibernate in woods and other
sheltered sites (see Alford et al., 2003 and literature
therein). With the exception of pollen beetles these pest
species have never been studied in a landscape context.
Because pest species must migrate to colonize the crop the
spatial patterns of OSR fields and potential overwintering
sites were assumed to affect their abundances.
The specific objectives of this study were to determine
(i) whether field and landscape characteristics influence
the major OSR pests in different ways, (ii) at which
spatial scales the influence of landscape variables are
effective and (iii) possible consequences for field and
landscape management. Generally, we hypothesized
that the abundances of these specialized pests should
be more related to variables describing OSR in the
landscape than to non-crop variables and that less
mobile D. brassicae would respond to landscape
characteristics at smaller scales than the more mobile
Ceutorhynchus and Meligethes species.
Materials and methods
Study area and landscape structure
The study region (about 240 km2) is located in the
relatively flat region of Lower Austria, about 40 km east
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684 J.G. Zaller et al. / Basic and Applied Ecology 9 (2008) 682–690
of Vienna (altitudes of arable land ranging from
130–250 m a.s.l.; coordinates of the central area:
161570 E, 481040 N). During the study period from April
to July 2005 the average temperature was 16.9 1C; the
rainfall during this period amounted to 336 mm (average
temperature and rainfall in these months between 1990
and 2004 was 16.0 1C and 230 mm, respectively; weather
data were provided from the meteorological station
Bruck/Leitha located about 16 km to the west of the
study region). The region mainly consists of farmland on
chernozem soils cropped according to integrated pest
management guidelines (main crops: wheat, maize,
barley, OSR, sunflower, sugar beet, poppy seed) inter-
spersed with semi-natural non-crop areas such as
fallows, hedges and forest remnants. The 29 OSR study
fields were randomly selected from this region that
covered a landscape complexity gradient ranging from
structurally poor to complex (average minimum distance
between study fields: 1198 m). Scale effects were inves-
tigated by calculating landscape variables at eight
circular sectors ranging from 250 to 2000 m radius
around our study plots using the software packages
ArcGis 9.1 and ArcView GIS 3.3 (ESRI Redlands, CA,
USA). Intensive field surveys using real-color orthopho-
tos (minimum resolution 0.25 m) were conducted in 2005
to assess the current landscape composition. Landscape
variables calculated were proportions of OSR fields,
non-crop area, area of grassy fallows and woody areas
(see Appendix A: Table 1). To characterize the landscape
diversity of the study area, the Shannon–Wiener index
(Krebs, 1994) was calculated based on 14 habitat types
(arable land, grassy fallows, woody fallows, copse, dry
grassland, dry shrubland, dry forests, riparian forest,
woods along rivers, hedges, settlements, aquatic areas,
roads, vineyards). Isolation of OSR fields in the land-
scape was calculated by using a negative exponential
weighting function, where isolation was calculated based
on the distances of neighbouring OSR fields to our study
field and OSR area of a given spatial scale using the
formula: Isolation OSRi ¼ "S(e"distance # OSR areaj)/
Se"distance (Kruess, 2003). Information on field variables
was obtained by a questionnaire among the participating
farmers (see Appendix A: Table 1). The soil index
specifies the natural yield capacity of a field in relation to
the highest yielding area of the country (0ysoil with
lowest yield capacity, 100ysoil with highest yield
capacity) and is an integrative measure of soil quality
that encompasses additionally the soil type, humus
content, soil depth, texture, density, structure, lime
content, gleying and soil congregation (ÖBG, 2001).
Crop management and pest sampling
OSR (cv. Californium) was sown by the farmers on
the 29 fields between 20 August and 14 September 2004
at seeding rates between 3.1 and 5.2 kg ha"1. OSR fields
were fertilized and treated with herbicides, fungicides
and insecticides following common agricultural practice
until December 2004. Starting January 2005, 1 ha of
each of the OSR fields was excluded from pesticide
applications and used for sampling insect pests and crop
plants for this study. A buffering zone of at least 10 m to
the next sprayed OSR was kept. The preceding crop was
winter barley for 18 fields, winter wheat for 10 fields and
poppy seed for one field (there was no relation between
the preceding crop and total pest abundances:
r2 ¼ 0.032, P ¼ 0.354; logistic regression analysis).
Stem weevil larvae were sampled in late April 2005 by
removing 25 randomly chosen OSR plants along a 50 m
transect located in the central area of each study field,
stems were dissected subsequently and weevil larvae
therein counted. Two stem weevils (C. pallidactylus
and C. napi) are major pests in the region with
C. pallidactylus comprising more than 80% of the stem
weevil abundance. Adult pollen beetles were sampled
during OSR flowering in late April 2005 at sunshine and
low wind velocity between 10 a.m. and 2 p.m. on 25
randomly chosen OSR plants along a 50 m transect
located in central area of each field. A plastic bag was
put over the top raceme, the raceme was cut off, and
pollen beetles were counted in the laboratory. Pollen
beetle communities comprise several species of the genus
Meligethes. In our study region, M. aeneus is the
predominant one, M. viridescens can be observed only
sporadically (Kraus & Kromp, 2002). Brassica pod
midge larvae (D. brassicae W) were assessed in late May
2005 in 100 randomly chosen pods from the top racemes
of OSR plants along a 50 m transect located in the
central area of each field after dissecting the pods in the
laboratory.
For better comparability of fields, all pest abundance
data were expressed on a 1-m2 basis by using OSR stand
density data (average number of stems assessed in two 1-
m2 frames). OSR plants in the study plots were cut
within two 1-m2 frames per field in the third week of
June 2005 and yields were assessed after threshing of
pods and wind sifting.
Statistical analyses and calculations
Effects of field and landscape variables on abundance
of OSR pest species were analysed using an ordinary
least-squares regression (OLS) model. Response vari-
ables were tested for normality using Shapiro–Wilk
W-statistic and log-transformed when necessary to meet
criteria for regression analyses (Zar, 1996). To analyse
scale effects, we compared the predictive power of
landscape variables measured at the eight spatial scales
(250, 500, 750, 1000, 1250, 1500 and 2000 m radius
centered in our study fields) in a univariate regression
 ARTICLE IN PRESS
J.G. Zaller et al. / Basic and Applied Ecology 9 (2008) 682–690
approach. Prior to multivariate analyses we tested the
predictor variables for intercorrelations (see Appendix
A: Table 2). Of the intercorrelating variables we
excluded those from the subsequent analyses that did
not characterize specific non-crop elements (N fertiliza-
tion level, landscape diversity, non-crop area, OSR
isolation). Significant predictor variables were selected
by performing a forward stepwise selection followed by
a backward elimination procedure (Yee & Mitchell,
1991). Possible non-linear effects were accounted for by
using restricted quadratic and cubic spline functions
(Harrel, 2001). Linearity of the predictor variables were
tested by comparing linear and non-linear terms (Wald
tests, F-statistics; Po0.05). At each step of the variable
selection, the significance of partial effects was tested by
dropping each predictor term from the model (Wald
tests, F-statistics; Po0.05). Because variables arising
from such nested measurement designs can be corre-
lated, we allowed only the most significant of a group of
correlated variables (e.g., one landscape variable at
different radii and different variables at the same radius)
to be included into the next step of variable selection. To
test for possible spatial autocorrelation we used Moran’s
I correlograms of residuals of the landscape regression
model (Legendre & Legendre, 1998; Lichstein, Simons,
Shriner, & Franzreb, 2002).
Results
The investigated landscape sectors varied consider-
ably in their structure and complexity, and OSR fields
differed in soil index and nitrogen fertilization (see
Appendix A: Table 1). Overall, we found great variation
in pest abundances among the studied OSR fields with
mean abundances (7SE) of brassica pod midge
126716 m"2, pollen beetle adults 346777 m"2 and stem
weevil larvae 3074 m"2. Abundance of pollen beetle
adults was significantly positively correlated with stem
weevil larvae (r ¼ 0.650, Po0.001) and pod midge
larvae (r ¼ 0.440, P ¼ 0.015); abundance of pod midge
larvae was not correlated with stem weevil larvae
(r ¼ 0.318, P ¼ 0.086).
Univariate regression analyses showed that only
abundances of pollen beetles (r2 ¼ 0.359, P ¼ 0.001)
and stem weevils (r2 ¼ 0.291, P ¼ 0.002) but not of pod
midges (r2 ¼ 0.015, P ¼ 0.492) were positively related to
soil index; nitrogen fertilization levels were not sig-
nificantly related to pest abundances. Pollen beetles
showed a negative relationship to OSR area (Fig. 1B)
and positive relations with the isolation of OSR in
the landscape (Fig. 1E), the proportion of non-crop
(Fig. 1K, woody areas (Fig. 1N) and landscape diversity
(Fig. 1H. Stem weevil larvae were negatively related to
OSR area (Fig. 1C), but positively related to OSR
685
isolation (Fig. 1F) and the proportion of woody areas
(Fig. 1O). Abundance of pod midge larvae was
positively related to landscape diversity (Fig. 1G) and
the proportion of woody areas (Fig. 1M).
For most pest species responses to landscape variables
showed clear maxima at certain scales (Fig. 1).
Abundance of pod midge larvae was explained by
proportion of woody areas up to 500 m only but no
relation to woody areas at greater radii (Fig. 1M).
Additionally, pod midge abundance was significantly
associated with landscape diversity at 1000 and 1250 m
radius but was unrelated at other scales (Fig. 1G).
Pollen beetle abundance was related to OSR area across
all tested scales (Fig. 1B) and related to OSR isolation at
a low (250 m) and larger radii (X1250 m; Fig. 1E).
Additionally, pollen beetle abundance was significantly
related to landscape diversity at 1500 m (Fig. 1H, non-
crop area between 1250–1750 m radius (Fig. 1K and
woody areas above a radius of 250 m (Fig. 1N).
Abundance of stem weevil larvae was significantly
explained by OSR area between 500 and 1000 m (Fig.
1C), by the isolation of OSR at 750 and 2000 m radius
(Figs. 1F) and by woody areas between 250 and 1000 m
radius and at 2000 m radius (Fig. 1O). Abundances of
tested pest species showed generally no relation to the
area of grassy fallows (data not shown).
Multivariate OLS analysis for pod midge abundances
revealed that no other variable except the proportion of
woody areas could significantly enhance the predictive
power of the model (univariate model result: r2 ¼ 0.279,
P ¼ 0.009). The multivariate model for the pollen
beetles (r2 ¼ 0.770, Po0.001) contains two variables,
the proportion of OSR at 1000 m radius (partial
r2 ¼ 0.368, Po0.001) and soil index (partial r2 ¼ 0.102,
P ¼ 0.0137) with a distinct negative response to OSR
area and a curvilinear relationship to soil index. The
plot of the partial effects indicated that pollen beetle
abundance was affected by the soil index mainly below
4% OSR area in the landscape (Fig. 2A). Above 4%
OSR area soil index played only a marginal role in
modulating the primary correlation of pollen beetle
abundance to OSR area. Maximum pollen beetle
abundance is predicted by this model at lowest levels
of OSR area and average soil index values (Fig. 2A).
The final model for the stem weevil data (r2 ¼ 0.550,
P ¼ 0.019) also consisted of two variables: soil index
(partial r2 ¼ 0.335, P ¼ 0.001) and proportion of woody
areas at a radius of 250 m (partial r2 ¼ 0.116,
P ¼ 0.017). The plot of the partial effects shows that
abundance was highest when soil index values were
slightly above average and proportion of woody areas is
high (Fig. 2B). While abundance showed a linear
response to woody area, the relation to soil index is
curvilinear and resembles a saturation curve (Fig. 2B).
The Moran’s I correlograms gave no indication for
spatial patterns in the residuals of the OLS model.
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686 J.G. Zaller et al. / Basic and Applied Ecology 9 (2008) 682–690

Fig. 1. Relationships between abundance of pod midge larvae (D. brassicae), adult pollen beetles (M. aeneus+M. viridescens) and
stem weevil larvae (C. pallidactylus+C. napi) and landscape variables derived from univariate ordinary least-square regression
analyses. Scale-dependent predictors are displayed for all investigated radii. Asterisks denote statistical significance: *Po0.05,
**Po0.01. With the exception of graphs B+C all relationships are positive.

OSR yield was significantly negatively correlated with
pollen beetle abundance (Fig. 3B), but unrelated to the
abundance of pod midge larvae (Fig. 3A) or stem
weevils (Fig. 3C).
Discussion
Pest responses to field characteristics
Abundances of pollen beetle adults and stem weevil
larvae showed strong relations to soil quality but
remained unrelated to nitrogen fertilization levels; pod
midges showed no relation to field characteristics tested.
Because pollen beetles and stem weevils do not over-
winter or pupate in current OSR fields, their soil quality
is unlikely to have any direct effect on their current
abundance. However, it is most likely that bottom-up
effects via the nutritional quality or the canopy
microclimate of OSR could have influenced the search-
ing efficiency of pollen beetles and stem weevils and thus
altered their abundances (Walters, Young, Kromp, &
Cox, 2003). In fact, glucosinolates and their catabolites
in OSR have been found to be important cues for host
selection by pests of crucifers, aiding both orientation to
and recognition of the host plant (Bartlet, 1996). In the
current study we used a double-zero cultivar that was
reduced in its glucosinolate and erucic acid content,
however other secondary compounds may have been
affected by soil quality (Nicholsorians, 1991). The
curvilinear shape of the relationship between pollen
beetle and stem weevil abundance and soil index with a
maximum at average levels indicates that at low soil
index, OSR quality was not suitable for pests. On the
other hand at a higher soil quality the crop could have
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J.G. Zaller et al. / Basic and Applied Ecology 9 (2008) 682–690
exhibited a better ability to protect itself from herbivores
via the production of secondary compounds (Cipollini &
Bergelson, 2002). It remains to be tested experimentally,
whether soil quality could also have affected the colour
and odour of OSR plants and thereby altered their
attractiveness to pest species.
Pest responses to landscape characteristics
Generally, we hypothesized our specialist pest species
to be more responsive to the proportion and isolation of
OSR than to non-OSR elements in the landscape (Holt,
Fig. 2. Partial effects of field and landscape variables on the
abundance of (A) pollen beetles adults (M. aeneus+M.
viridescens) and (B) stem weevil larvae (C. pallidactylus+C.
napi) derived from multivariate ordinary least-square regres-
sion analyses.
Fig. 3. Relationship between OSR yield and abundance of pod midge larvae (D. brassicae), pollen beetles (M. aeneus+M.
viridescens) and stem weevil larvae (C. pallidactylus+C. napi). Formulae, r2 and P-values derived from linear regression fits (n ¼ 29).
687
Lawton, Polis, & Martinez, 1999). This was confirmed
for pollen beetles, partly confirmed for stem weevils but
not for pod midges. Pollen beetles and stem weevils were
more abundant in landscapes with less OSR area and
less abundant when more OSR area was available. This
is in contrast to findings by Thies, Steffan-Dewenter,
and Tscharntke (2003), however could be explained by
the different methodological approaches. The coloniza-
tion by pollen beetle of a few potted OSR plants located
in old field margins in the former study may have
differed considerably from that of field-grown plants in
the current study. Generally, the pest distribution
patterns could be expected to mirror areas of OSR
and overwintering sites of the preceding year that
enabled the buildup of a certain landscape pest pool
that was then dispersed among available OSR area in
the current year (Hokkanen, 2000). However, since in
our region OSR cropping history and non-crop areas
did not change considerably during the last years
because agri-environmental programmes provided sub-
sidies to promote both OSR cropping and extensifica-
tion in the region (BMLFUW, 2006), we assume that
pest migration processes are most likely responsible for
these findings.
Pollen beetle abundance and non-crop area were
positively related suggesting that more complex land-
scapes support a greater variety of alternative host
plants for pollen beetles, and thereby complex land-
scapes may have enhanced pest populations (Frenzel &
Brandl, 1998). However, this finding appears to be in
contrast to that of others (Thies et al., 2003; Thies &
Tscharntke, 1999) who showed a negative relationship
between pollen beetle damage (i.e., number of podless
stalks) and non-crop area. Because pollen beetle
abundance and damage caused are well correlated
(Zaller, Moser, Drapela, Schmöger, & Frank, 2008)
several explanations could be considered for this
discrepancy. First, the parameter non-crop area per se
does not adequately describe landscapes because non-
crop usually encompasses various habitats (e.g., fallows,
woody areas, meadows) that can exert contrasting
effects on pest abundances. For instance, in our study
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688 J.G. Zaller et al. / Basic and Applied Ecology 9 (2008) 682–690
non-crop was mainly comprised of grassy fallows and
woody areas, but only the latter showed a significant
effect on pest abundances. Moreover, in our study
grassland elements were mainly composed of grassy
fallows while in the former studies pastures and
meadows formed the grassland elements (Thies et al.,
2003; Thies & Tscharntke, 1999). Second, contrasting
findings might again result from the fundamentally
different methodological approaches used. The positive
relations between pollen beetles and non-crop in our
study also suggest that natural enemies promoted by
non-crop areas (e.g. hymenopterous parasitoids – Thies
& Tscharntke, 1999; carabid beetles – Östman et al.,
2001b; spiders – Schmidt, Roschewitz, Thies, &
Tscharntke, 2005; Drapela et al., unpublished data)
had no detrimental influence on pests in our region.
Indeed we observed that parasitoids of pollen beetles
seemed to play only a minor role in our landscapes/
region (Zaller et al., unpublished data) and there is
evidence that besides beneficial organisms also pests are
supported by non-crop structures (Thies, Roschewitz, &
Tscharntke, 2005).
Pod midge abundance was unrelated to OSR vari-
ables but positively related to landscape diversity and
woody areas. This finding seems somewhat surprising
because pod midge adults are reported to emerge from
the OSR field of the last year and in the following spring
migrate to the current OSR fields (Alford et al., 2003).
However, it also has been observed that second and
third generations of these polyvoltine pests are over-
wintering in stands of other Brassicaceae (Paul, 2003).
Thus, our data suggest that for this species a more
diverse landscape with more woody areas increases the
likelihood of finding alternative host plants. However,
admittedly our understanding of landscape effects on
pod midges is still too scarce to be able to further
interpret our findings.
Two general trends for the three species groups could
be seen in our data: (i) pest abundances were consis-
tently positively related to woody areas, and (ii) an
absence of any relation between pest abundances and
grassy fallows. For pollen beetles, the positive effect of
woody areas can be explained by their ability to serve as
overwintering sites. For stem weevils and pod midge
that are thought to overwinter in non-woody areas it is
possible that woody areas in the vicinity of the actual
overwintering habitats could have climatically influ-
enced these habitats (e.g., reduced wind exposition or
increased relative humidity; Foggo, Ozanne, Speight, &
Hambler, 2001). No detrimental effects of grassy fallows
on populations of OSR pests could be identified
although these sites have frequently been shown to be
important overwintering habitats for natural enemies
of OSR pest species (Östman et al., 2001b; Thies
et al., 2003). The lack of a relation between pests and
grassy fallows also suggests that a widely used assump-
tion that fallows are refuges for insect pests (Lethmayer,
Nentwig, & Frank, 1997) could not be substantiated by
our data.
Spatial aspects of pest responses
Our data showed that the scale at which each insect
group is influenced is related to each groups dispersal
capacity. Generally, pod midges seemed to respond to
landscape factors at small (250–500 m; woody areas) and
medium scales (1000–1250 m; landscape diversity),
pollen beetles showed effects at medium to large scales
(1000–2000 m; all variables except grassy fallows) while
stem weevils were related to landscape factors mainly at
small to medium scales (500–1000 m; OSR area, isola-
tion OSR, woody areas). A limitation of available OSR
food at certain scales seemed unlikely to be responsible
for these relationships, however dispersal limitations
might be the reason for these patterns (Kruess &
Tscharntke, 1994, 2000; Tscharntke & Kruess, 1999;
With, Cadaret, & Davis, 1999). Pod midges are known
to be weak fliers with females usually dispersing no more
than a few hundred meters from their emergence sites.
This might explain their response to woody areas at the
very smallest scales, however our data also indicate that
responses at medium scales are possible. In contrast,
pollen beetles can easily cross distances greater than that
considered in the current study (Fritzsche, 1957) and are
therefore affected by landscape characteristics on
medium to large scales. Information on the mobility of
stem weevils is generally scarce but they are reported to
preferably make short flights (Schmutterer, 1956). Based
on the current results, however, this did not seem to
hinder them to react to landscape effects across all
scales.
Overall, this study is a first attempt to understand
how abundances of different pest species might be
affected by landscape characteristics and at what spatial
scale this occurs, however, much more research is
necessary to be able to elucidate the underlying
processes particularly regarding pest migration patterns
and dispersion between metapopulations of pests in a
landscape. From a farmer’s perspective pollen beetles
appear to be the most important species because they
showed the most pronounced negative impact on yields,
however, data also showed that the studied pest species
were generally co-occurring in our fields. Our results
suggest that crop rotation schemes commonly in place in
order to reduce the carry-over of OSR pest and disease
problems should be expanded by a landscape perspec-
tive. However, more multi-scale assessments conducted
in different regions are necessary to adequately pinpoint
how habitat features and the spatial configuration of
crop and non-crop elements can affect the abundance of
pest populations.
 ARTICLE IN PRESS
J.G. Zaller et al. / Basic and Applied Ecology 9 (2008) 682–690
Acknowledgments
We are grateful to the farmers for participating in this
research project, the staff of the University Research
Farm Groß-Enzersdorf for providing post-harvest
equipment and to the Austrian Science Fund for
supporting J.G.Z., D.M. and T.D. (Grant no. P16972).
The regional governments of Lower Austria and Burgen-
land provided maps and aerial photographs of the project
area. Help by Norbert Schuller and Erhard Tesarik in the
field and laboratory is gratefully acknowledged.
Appendix A. Supplementary materials
Supplementary data associated with this article can
be found in the online version at doi:10.1016/j.baae.
2007.10.004.
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