ARTICLE IN PRESS

Basic and Applied Ecology 10 (2009) 208–215 www.elsevier.de/baae

Interacting effects of wind direction and resource distribution

on insect pest densities
Dietmar Mosera,b,!, Thomas Drapelaa, Johann G. Zallera, Thomas Franka
a
Institute of Zoology, Department of Integrative Biology and Biodiversity Research, University of Natural Resources and Applied
Life Sciences Vienna, Gregor Mendel Strasse 33, A-1180 Vienna, Austria
b
Vienna Institute for Nature Conservation and Analyses, A-1090 Vienna, Austria

Abstract
Considerable effort has been made to investigate how landscape composition and spatial structures of habitats
influence distribution patterns of species. In particular, specialist insect herbivores are known to be affected by spatial
and temporal accessibility of their host plants. We studied three important insect pests of oilseed rape (OSR):
Meligethes aeneus, Ceutorhynchus pallidactylus and Dasineura brassicae. In a landscape with northwest winds
prevailing, we analysed their densities by comparing the predictive power of OSR area in differently orientated
landscape sectors. Regression analyses showed that OSR area downwind from a sample site explained up to 72% of
the variance in the density of M. aeneus, whereas OSR area in other directions had little effect. The correlation between
downwind OSR area and M. aeneus density was negative and observable up to a distance of 1250 m. In contrast, the
densities of C. pallidactylus and D. brassicae showed little response to OSR area in whatever direction. We suggest that
mainly resource detection mechanisms and dispersal capabilities are responsible for the detected patterns: Odour-
orientated upwind anemotaxis apparently drives directional dispersal of mobile species (M. aeneus). However, OSR
area along the dispersal path seems to reduce pest density in upwind direction, because the majority of individuals
detect resource patches early during the dispersal process. For less mobile species (C. pallidactylus and D. brassicae),
similar effects were not detectable at the landscape scale because dispersal capabilities probably were too short.
r 2008 Gesellschaft für Ökologie. Published by Elsevier GmbH. All rights reserved.

Zusammenfassung
Eine der Grundfragen der landschaftsökologischen Forschung ist die nach dem Einfluss der Landschaftszusam-
mensetzung und der räumlichen Struktur von Habitaten auf das Vorkommen von Arten. Vor allem für spezialisierte
herbivore Insekten ist die räumliche und zeitliche Erreichbarkeit ihrer Wirtspflanzen von entscheidender Bedeutung. In
dieser Studie untersuchten wir inwieweit die Abundanzen dreier wichtiger Rapsschädlinge (Meligethes aeneus,
Ceutorhynchus pallidactylus und Dasineura brassicae) in einer Landschaft mit vorherrschend nordwestlicher
Windrichtung von in unterschiedlichen Himmelsrichtungen gelegenen Rapsanbauflächen abhängig ist. Die
Regressionsanalysen ergaben dass M. aeneus eine starke Abhängigkeit zur Rapsfläche auf der windabgewandten
Seite zeigte (R2 ¼ 0,72), während die windzugewandte Seite keine signifikante Rolle spielte. Der Zusammenhang
zwischen M. aeneus-Abundanz und Rapsflächenanteil in windabgewandten Landschaftsausschnitten war negativ und

!Corresponding author. Current address: Institute of Zoology, Department of Integrative Biology and Biodiversity Research, University of
Natural Resources and Applied Life Sciences Vienna, Gregor Mendel Strasse 33, A-1180 Vienna, Austria. Tel.: +43 1 47654 3205;
fax: +43 1 47654 3203.
E-mail address: dietmar.moser@vinca.at (D. Moser).

1439-1791/$ - see front matter r 2008 Gesellschaft für Ökologie. Published by Elsevier GmbH. All rights reserved.
doi:10.1016/j.baae.2008.03.008
 ARTICLE IN PRESS
D. Moser et al. / Basic and Applied Ecology 10 (2009) 208–215 209

beobachtbar bis zu einer Distanz von 1250 m. Im Gegensatz zu M. aeneus zeigten C. pallidactylus and D. brassicae
kaum eine Reaktion auf Rapsflächenanteilen in unterschiedlichen Richtungen. Wir vermuten, dass vor allem
Nahrungssuchstrategien und Ausbreitungsvermögen der untersuchten Arten über die Stärke des Zusammenhangs
entscheiden. Geruchsinduzierte windwärts gerichtete Suchflüge bewirken eine gerichtete Ausbreitung von mobilen
Arten (M. aeneus). Eine hohe Dichte an Rapsfeldern entlang des Ausbreitungsweges scheinen die Abundanzen von M.
aeneus in der windzugewandten Seite zu reduzieren, da ein Großteil der Individuen früh ein Rapsfeld entdeckt und den
Suchflug beendet. Geringeres Ausbreitungsvermögen dürfte für das Fehlen derartiger Zusammenhänge auf
Landschaftsebene bei den weniger mobilen Arten C. pallidactylus und D. brassicae verantwortlich sein.
r 2008 Gesellschaft für Ökologie. Published by Elsevier GmbH. All rights reserved.

Keywords: Brassica napus; Oilseed rape; Meligethes aeneus; Ceutorhynchus pallidactylus; Dasineura brassicae; Density:area relation;
Dispersal behaviour; Landscape ecology; Olfactory search; Wind dispersal

Introduction known about the perceptual range of the species

Knowledge of dispersal behaviour is crucial for
understanding distribution patterns of various impor-
tant pest species. Understanding the underlying princi-
ples and the spatial range of dispersal processes will
improve our ability to predict pest emergence and will
help to develop appropriate habitat management
strategies to reduce herbivore pressure.
In the current investigation, we examined three
important oilseed rape (OSR) pest insects: the pollen
beetle (Meligethes aeneus), the cabbage stem weevil
(Ceutorhynchus pallidactylus) and the brassica pod
midge (Dasineura brassica). In a previous study (Zaller,
Moser, Drapela, Schmöger, & Frank, 2007), we
observed that M. aeneus density was strongly negatively
correlated with OSR area in the surrounding of our test
sites, whereas C. pallidactylus density showed a much
lower dependency, and D. brassicae proved to be
uncorrelated with OSR area. Dispersal processes and
host detection mechanisms are assumed to be key
factors controlling relationships between the area of
resource habitat and the density of insect populations
(Bukovinszky, Potting, Clough, van Lenteren, & Vet,
2005; Englund & Hambäck, 2007; Hambäck &
Englund, 2005; Hambäck, Summerville, Steffan-Dewen-
ter, Krauss, Englund et al., 2007; Otway, Hector, &
Lawton, 2005). Generally, visual searchers are assumed
to show a negative density:area relationship, whereas the
density of olfactory searchers should respond positively
to resource area (Bukovinszky et al., 2005; Hambäck &
Englund, 2005; Hambäck et al., 2007). Several olfact-
ometer and field-trapping studies have demonstrated
that both visual and olfactory cues (isothiocyanates) are
important for the location of host plants by M. aeneus
(Evans & Allenwilliams, 1994; Jonsson, Rosdahl, &
Anderson, 2007; Smart & Blight, 2000; Williams,
Frearson, Barari, & McCartney, 2007), Ceutorhynchus
spp. (Bartlet, Blight, Lane, & Williams, 1997; Evans &
Allenwilliams, 1993) and D. brassicae (Murchie, Smart,
& Williams, 1997). While the attraction of pest species
by visual and olfactory cues is well documented, little is
(Schooley & Wiens, 2003; Williams et al., 2007; Zollner
& Lima, 1999). Olfactometer experiments and mark-
release-recapture studies with odour-baited traps (Cook,
Bartlet, Murray, & Williams, 2002; Evans & Allenwil-
liams, 1994) typically work at fine scales (within a metre
and 20 m, respectively), while field trap experiments do
not provide information on the origin of the insects
trapped (Williams et al., 2007). Wind direction and
upwind migration might play an important role in the
dispersal of pest species (Ferguson, Barari, Warner,
Campbell, Smith, Watts et al., 2006; Williams et al.,
2007; for other species: Meats & Hartland, 1999;
Schooley & Wiens, 2003) but there is little information
on the spatial scales at which these factors operate (e.g.
20 m for M. aeneus in Evans & Allenwilliams, 1994).
Since our study region is characterised by prevailing
northwest winds, we assume that wind will be an
important factor controlling distribution patterns of
pest species. For the current study, we hypothesised that
both wind direction and OSR area available in the
landscape interactively influence pest species densities in
OSR fields. This hypothesis was tested by analysing the
predictive power of the OSR area in surrounding
landscape segments in different directions. If wind
played any role, there should be considerable differences
in the effect of OSR area on pest species density,
depending on its location relative to the point of
observation, up- or downwind.
Methods
Study area
The study region is one of the most important OSR
growing areas in eastern Austria and is located about
40 km east of the city of Vienna near the Hungarian
border, south of the river Danube (LL: 48110 N 161520 E;
UR: 48180 N, 17150 E, about 240 km2). The area mainly
consists of intensively managed farmland on chernozem
 ARTICLE IN PRESS
210 D. Moser et al. / Basic and Applied Ecology 10 (2009) 208–215
soils with a varying portion of different-aged fallows,
hedges and forest remnants (average landscape compo-
sition: crop: 67.3%; woody areas including forests,
hedges and scrubland: 12%; other non-crop: 16.3%;
settlements: 4.4%). The main crops are wheat, maize,
barley, OSR, sunflower, sugar beet, poppy seed and
vineyards.
Species
M. aeneus (pollen beetle) is univoltine. Adults emerge
in spring and invade OSR fields where they lay their eggs
in flower buds. Although the pollen beetle is polypha-
gous and feeds on pollen of several plants, oviposition
occurs on species of the family Brassicaceae exclusively.
After 19–35 days, larvae leave the flowers and pupate in
the soil where new adults appear after 10–18 days. After
emergence, adult beetles feed on pollen from many
families before they hibernate in woods and other
sheltered habitats (Alford, Nilsson, & Ulber, 2003; Paul,
2003). Pollen beetles are known to have a large dispersal
range with flight distances between 1 and 3 km day"1
(Stechmann & Schütte, 1976).
C. pallidactylus (cabbage stem weevil) is also uni-
voltine and adult beetles migrate to OSR fields in spring
from bud stages onwards and oviposit on the underside
of petioles where the larvae tunnel into the stems. After
3–5 weeks, larvae vacate the host plant and pupate in
the soil. After hatching in summer, adults feed on leaves
of other species of the family Brassicaceae before
hibernation (Alford et al., 2003; Paul, 2003). We are
not aware of any study providing clear information on
overwintering habitats and dispersal range.
D. brassicae (brassica pod midge) has two to three
generations per year. On emergence from hibernation,
adults migrate to OSR fields where they oviposit in
pods. Larvae pupate in the soil of the OSR field, where a
small proportion immediately starts a diapause until the
next spring. The others hatch for a second and some-
times a third generation, which use alternative crucifer-
ous species as hosts. The pod midge is assumed to be a
weak flyer, dispersing no more than a few hundred
metres from its emergence site (Alford et al., 2003; Paul,
2003).
Species data
Pest species densities were determined from samples
collected on 29 randomly selected OSR plots (1 ha).
OSR (Brassica napus, cv. Californium) was sown by the
farmers between 20 August and 14 September 2004
(average seeding rate: 4.2 kg ha"1). Fertilizer, fungicide,
herbicide and insecticide application was permitted until
December 2004, following common farming practice.
Samples were collected keeping a distance of at least
10 m to the next sprayed field. Adult pollen beetles were
sampled from 25 randomly selected top racemes per
sampling plot on 27 April 2005. Racemes were stored in
plastic bags and beetles were counted in the laboratory.
The stem weevil larvae were assessed from 25 randomly
selected OSR plants per sampling plot on 27 April 2005.
The stems were dissected and inspected for weevils. Data
on pod midge larvae were obtained from 100 randomly
chosen pods from top racemes per sampling plot on
23 May 2005. Insect densities were calculated as the
number of individuals m"2, by using OSR stand density
data (average number of plants assessed in two 1 m2
frames).
Assessing wind direction and landscape data
Data on wind direction were obtained from a nearby
windfarm. We used measurements of seven windmills
(all located within our study area) taken at 10 min
intervals (February–May 2005) to calculate the average
proportion of wind from different directions. A detailed
land-cover map was drawn up based on intensive field
surveys using real colour orthophotos (resolution
0.25 m) in 2005. The assessment comprised all land-use
and habitat-types within a buffer radius of 2000 m
around each sampling plot. OSR area was calculated as
follows: We constructed 901 sectors of circles with radii
of 250, 1250 and 1750 m, centred in the midpoint of our
sampling plots. The 1250 and 1750 m sectors were
designed as donut sectors with inner radii of 250 and
1250 m, respectively. The area was 4.91 ha for the 250 m
sectors and 117.81 ha for the 1250 and 1750 m sectors.
Fig. 1 shows the SE sector as an example, but sectors
were constructed for all cardinal and intercardinal
directions. Areal percentages of OSR were then calcu-
lated upon intersection with the land cover map. OSR
area within a given sector ranged from 0.002 to 4.88 ha
(0.7070.90; mean 71 SE) for the 250 m sector, from 0
to 38.78 ha (6.3475.52) for the 1250 m sectors and
0–35.80 ha (5.6375.28) for the 1750 m sectors. Overall,
the area of single OSR fields ranges from 0.11 to
18.87 ha (2.2472.55). We used ARC-GIS 9.1 and
ArcView GIS 3.3 (ESRI Redlands, CA, USA) for
analysing geographical data.
Statistical analyses
We examined the mean effect of OSR area on the
density (individuals m"2) of pollen beetles, stem weevils
and pod midges by means of 24 (eight directional sectors
# three radii) ordinary least-squares regression (OLS)
models. The response variables were tested for normal-
ity by the Shapiro–Wilk W-statistics (Zar, 1996) and log
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D. Moser et al. / Basic and Applied Ecology 10 (2009) 208–215
Fig. 1. Schematic example for the construction of the land-
scape sectors (example for the SE sector). Each sector was
subdivided into an inner sector (radius: 250 m), an inner donut
sector (250–1250 m) and an outer donut sector (1250–1750 m).
Sectors were constructed for all cardinal and intercardinal
directions. OSR fields are marked in grey or black.
transformed if necessary. F-tests for comparison of
nested models were used to test for a significant effect of
the predictor variables as compared to intercept-only
models. Non-linearity was tested by using restricted
cubic spline functions with three knots (Harrel, 2001).
Non-linearity of the predictor variables was tested by
comparing linear and non-linear models with F-tests for
nested models. To test for short and long distance effects
of OSR area, we designed multivariate OLS models by
successively adding the terms of all three radii and
testing for significant model improvement (F-tests).
Interaction terms were tested, too. ANOVA analyses
were used to test whether direction had any effect on the
OSR area. All statistical analyses were performed with
S-PLUS 7.0 for Windows (Insightful Corporation,
Seattle, WA, USA).
Results
Pest density varied considerably among the investi-
gated OSR fields with pollen beetle density ranging from
34 to 1798 individuals m"2, stem weevil larvae from 2 to
118 individuals m"2 and brassica pod midge larvae from
14 to 490 individuals m"2.
ANOVA analyses showed that direction generally
had no effect on the OSR area, with one exception for
the 250 m sectors, where the E and SW sectors had
significantly different areas of OSR fields. The analysis
of the windmill data confirmed prevailing NW winds in
our study region for the main dispersal period of the
investigated species (Fig. 2).
211
Fig. 2. Time proportion (%) of winds from different directions
in the study region (February–May 2005).
Pollen beetle density was significantly negatively
affected by OSR area in the short to medium distance
environments (0–250 and 250–1250 m radii, Fig. 3).
However, the effect of OSR area strongly depended on
direction relative to the sampling plots: for the short
distance radii the SE sector explained 62% (F ¼ 21.13
on 2 and 26 degrees of freedom, Po0.0001) of the
variance in the density data, whereas OSR area in the
NW sector was unrelated (Fig. 4, see Appendix A).
A similar pattern was observed for the 250–1250 m
sectors, where the predictive power of the SE sector
(R2 ¼ 0.51, F ¼ 14.40 on 1 and 27 degrees of freedom,
P ¼ 0.0001) was more than twice as that of the NW
sector (R2 ¼ 0.22, F ¼ 3.72 on 1 and 27 degrees of
freedom, P ¼ 0.0378). This pattern disappeared for the
sectors in the outer ring, which showed no clear trend
for any direction.
For the SE sector, OSR area at medium distances
(250–1250 m radii) had a significant partial effect on
pollen beetle density when added to a model with OSR
area at short distances (0–250 m radii) as predictor
(multiple R2 ¼ 0.72, F ¼ 22.04 on 3 and 25 degrees
of freedom, P ¼ o 0.0001). The outermost sector (SE
1250–1750 m) did not improve the model significantly.
No interaction terms were significant.
The results for the stem weevil and pod midge (Fig. 4,
Appendix A) did not show the clear trend that was
observed for the pollen beetle, but the response to OSR
area was also negative. At short distances (0–250 m
sectors) the stem weevils showed a weak negative
reaction to the SW sector and the pod midges to the S
sector. The results for the 250–1250 m sectors were more
similar to those of the pollen beetle. The predictive
power of OSR area was once again highest for the SE
sector, for both the stem weevils and the pod midges.
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212 D. Moser et al. / Basic and Applied Ecology 10 (2009) 208–215

Fig. 3. Scatter plot of pollen beetle density (individuals m"2) and OSR area for two SE sectors (0–250 and 1250–1250 m).

Fig. 4. Results of the regression analyses (R2) of the relationship between OSR area in eight 901 sectors and pest species abundance.
Sectors were sub-divided into three distance classes (0–250, 250–1250 and 1250–1750 m). Significant effects are marked by filled
symbols, insignificant effects by white symbols. Exact values of R2 are listed in Appendix A. Pollen beetle: circles, stem weevils:
squares, pod midge: triangles.

The analyses of the outer sectors (1250–1750 m) showed
highest R2 for the W and NW direction for the stem
weevils, whereas the pod midges revealed a weak but
significant reaction to the E sector.
Discussion
Our results showed that wind direction and OSR area
specifically affected densities of the three pest species.
The response of pollen beetle density to OSR area
showed a distinct polar pattern suggesting that beetle
density was predominantly affected by resource avail-
ability in the downwind landscape, whereas the upwind
landscape seemed to be of marginal importance (Figs. 2
and 4). This indicates that wind direction and resource
distribution interactively affected the searching beha-
viour of pollen beetles and confirms findings that they
use upwind anemotaxis to locate OSR fields in spring
(Evans & Allenwilliams, 1994, Williams et al., 2007).
Prevailing winds and odour-induced upwind anemotaxis
drive directional dispersal of mobile species that use
olfactory cues for resource location (Vickers, 2000),
because resource patches in upwind position have a
higher probability of being detected than resource
patches located downwind. One main process that
determines the abundance of pollen beetles in OSR
fields is the annual dispersion from overwintering to
feeding and reproduction sites. If we assume that this
dispersal process is mainly guided by anemotactic search
behaviour, the bulk of individuals will disperse upwind
and touch down as soon as they encounter an OSR field
(Fig. 5, cf. Hein, Pfenning, Hovestadt, & Poethke, 2004).
The density of individuals at a given site will, hence,
depend on the OSR area along the line between the
respective site and the overwintering habitat: the more
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D. Moser et al. / Basic and Applied Ecology 10 (2009) 208–215
Fig. 5. Diagram of a directional dispersal process (i.e. upwind
anemotaxis) of herbivorous insects from overwintering sites
(grey) to feeding and reproduction sites (white rectangles).
Dispersing beetles are symbolised by black arrows (the
thickness of the arrows indicates the number of beetles). The
abundance of beetles in the feeding and reproduction habitats
depends on the amount of resource habitats along the dispersal
path.
OSR area there is along this dispersal path the more
likely an individual will find an appropriate feeding and
reproduction habitat before reaching the sampling site.
Pollen beetle density is primarily related to immigra-
tion rates, decoupled from local reproduction. Immigra-
tion rate and resource area have been shown to be
inversely related, if patches are detected from a long
distance or if search is approximately linear (Englund &
Hambäck, 2007, Hambäck & Englund, 2005, Hambäck
et al., 2007). Negative density:area relations were
associated with visual searchers (e.g. butterflies), while
olfactory searchers (e.g. moths) should response posi-
tively (Hambäck et al., 2007). This seems to be in contrast
to our current results, if we assume that olfactory search
was the primary search strategy of pollen beetles.
However, predominant winds from one direction drive
directional dispersal of our species and unidirectional
transportation of odour particles may increase the
perceptual range (Vickers, 2000) suggesting that olfactory
cues gain importance for large range host detection in
regions with predominant winds from one direction.
Moreover, since pollen beetles are diurnal, localisation of
the host plant patches is aided by visual cues (Blight &
Smart, 1999; Evans & Allenwilliams, 1994; Giamoustaris
& Mithen, 1996; Ruther & Thiemann, 1997). However,
based on the current data we are not able to quantify the
contribution and perceptual range of visual versus
213
olfactorial orientation for large range host plant detec-
tion. Although the hypotheses of Englund and Hambäck
(2007) and Hambäck et al. (2007) provide appealing
explanations for our results, it has to be noted that our
analyses addressed area proportions, which is close to,
but not directly linked to patch area.
The density of pollen beetles at a sampling point will
certainly be co-determined by other factors, most
pronouncedly by the distribution of overwintering sites
in the landscape and the distribution of OSR fields in
the previous year. These two factors together control the
number of individuals that disperse in spring in the
‘‘catchment area’’ of a certain sampling point. They may
confound the relationship between pollen beetle density
and OSR area if they are not independent of OSR area
themselves. However, there is no indication of such
correlation, at least for overwintering sites, as OSR area
and area of woody habitats were generally uncorrelated.
Unfortunately, data on OSR field distribution in 2004
were not available.
The pronounced negative correlation between pollen
beetle and OSR area in south-eastern sectors could be
observed up to the 250–1250 m sectors, above that
distance the effect vanished. This suggests that the
perceptual range of the species must be somewhat
between 0 and about 1250 m, which is within the lower
range of the mean per day flight distance of 1–3 km
(Stechmann & Schütte, 1976).
In contrast to pollen beetle, stem weevil and pod
midge density was less strongly correlated with OSR area
in downwind directions. Although we also found a
higher R2 for SE sectors, at least for the 250–1250 m
radius, this pattern was much less pronounced. Lower
mobility of the pod midges (Alford et al., 2003; Paul,
2003) might be partly responsible for the weaker
response. Dissemination of the overwintering population
should, hence, be more local, and landscape-scale effects
are likely to be of minor importance. Interpreting results
for the stem weevil is hampered by a lack of autecolo-
gical data concerning hibernation habitats and dispersal
capabilities (Alford et al., 2003; Paul, 2003). Similarity
among patterns suggests that, like for the pod midges,
limited dispersal capabilities and/or perceptual range
might be responsible for the weaker correlation with
OSR area. Besides lower active dispersal capabilities,
higher rates of passive downwind dispersal may also
confound effects caused by active foraging. Wind speed
may not only affect perceptual range of an insect but also
its ability to fly upwind (Brady, Griffiths, & Paynter,
1995). Low wind speeds allow for active flight, while
stronger wind velocity may trigger passive dispersal.
While the preponderance of significant results in the SE-
directions for the inner circle and the inner doughnut
ring indicate active dispersal and response to host plant
cues, the shift to the NW direction in the outer ring
indicates the importance of passive flight in long distance
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214 D. Moser et al. / Basic and Applied Ecology 10 (2009) 208–215
dispersal. Unfortunately, there are no studies providing
reliable information allowing to differentiate between
active and passive dispersal in the studied species.
Perhaps, the weaker response of stem weevils and pod
midges might also be an effect of different sampling
methods used: while the pollen beetle data represented
adults, which were directly associated with immigration
rates, the weevil and pod midge data were counts of
larvae, which are not only driven by adult immigra-
tion but also by factors that control the subsequent
reproductive success. Finally, it should be mentioned
that, over the long run, the amount of OSR fields
enhances population size of pest species if we focus
on the whole landscape (Hokkanen, 2000). Thus, if
we compare different landscapes on a regional scale
there should be a positive correlation between species
abundance and resource availability (i.e. OSR area) so
our results will hold for within-landscape distribution
patterns only.
Our analyses suggest that interactions between wind
direction and resource distribution result in a negative
density:area relationship of olfactorily orientated OSR
pest species. Species density at a certain location is
probably more affected by resource distribution along
the dispersal path than by site area itself. However,
it remains to be investigated whether the patterns
described on a landscape scale for a highly mobile
species like the pollen beetle also hold for less
mobile species on a much smaller scale. In any case,
as our study is based on a pattern analysis only,
confirmation of our conclusions by direct mark-recap-
ture experiments will be necessary (e.g. Östrand &
Anderbrant, 2003).
Acknowledgements
We are grateful to the farmers for participating, to the
regional governments of Lower Austria and Burgenland
for providing maps and aerial photographs of the project
area. The EVN gratefully provided data on wind direc-
tions. Help by Norbert Schuller and Erhard Tesarik in the
field and laboratory is gratefully acknowledged. For their
comments on the manuscript, we thank Stefan Dullinger
and Barbara Holzinger. Peter Hambäck, Klaus Höve-
meyer and an anonymous referee gave valuable comments
on the manuscript. This study was financially supported by
the Austrian Science Fund (Grant no. P16972).
Appendix A. Supplementary Materials
Supplementary data associated with this article can
be found in the online version at doi:10.1016/j.baae.
2008.03.008.
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