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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
soils with a varying portion of different-aged fallows, 10 m to the next sprayed field. Adult pollen beetles were
hedges and forest remnants (average landscape compo- sampled from 25 randomly selected top racemes per
sition: crop: 67.3%; woody areas including forests, sampling plot on 27 April 2005. Racemes were stored in
hedges and scrubland: 12%; other non-crop: 16.3%; plastic bags and beetles were counted in the laboratory.
settlements: 4.4%). The main crops are wheat, maize, The stem weevil larvae were assessed from 25 randomly
barley, OSR, sunflower, sugar beet, poppy seed and selected OSR plants per sampling plot on 27 April 2005.
vineyards. 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
Species
23 May 2005. Insect densities were calculated as the
number of individuals m"2, by using OSR stand density
M. aeneus (pollen beetle) is univoltine. Adults emerge
data (average number of plants assessed in two 1 m2
in spring and invade OSR fields where they lay their eggs
frames).
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 Assessing wind direction and landscape data
the soil where new adults appear after 10–18 days. After
emergence, adult beetles feed on pollen from many Data on wind direction were obtained from a nearby
families before they hibernate in woods and other windfarm. We used measurements of seven windmills
sheltered habitats (Alford, Nilsson, & Ulber, 2003; Paul, (all located within our study area) taken at 10 min
2003). Pollen beetles are known to have a large dispersal intervals (February–May 2005) to calculate the average
range with flight distances between 1 and 3 km day"1 proportion of wind from different directions. A detailed
(Stechmann & Schütte, 1976). land-cover map was drawn up based on intensive field
C. pallidactylus (cabbage stem weevil) is also uni- surveys using real colour orthophotos (resolution
voltine and adult beetles migrate to OSR fields in spring 0.25 m) in 2005. The assessment comprised all land-use
from bud stages onwards and oviposit on the underside and habitat-types within a buffer radius of 2000 m
of petioles where the larvae tunnel into the stems. After around each sampling plot. OSR area was calculated as
3–5 weeks, larvae vacate the host plant and pupate in follows: We constructed 901 sectors of circles with radii
the soil. After hatching in summer, adults feed on leaves of 250, 1250 and 1750 m, centred in the midpoint of our
of other species of the family Brassicaceae before sampling plots. The 1250 and 1750 m sectors were
hibernation (Alford et al., 2003; Paul, 2003). We are designed as donut sectors with inner radii of 250 and
not aware of any study providing clear information on 1250 m, respectively. The area was 4.91 ha for the 250 m
overwintering habitats and dispersal range. sectors and 117.81 ha for the 1250 and 1750 m sectors.
D. brassicae (brassica pod midge) has two to three Fig. 1 shows the SE sector as an example, but sectors
generations per year. On emergence from hibernation, were constructed for all cardinal and intercardinal
adults migrate to OSR fields where they oviposit in directions. Areal percentages of OSR were then calcu-
pods. Larvae pupate in the soil of the OSR field, where a lated upon intersection with the land cover map. OSR
small proportion immediately starts a diapause until the area within a given sector ranged from 0.002 to 4.88 ha
next spring. The others hatch for a second and some- (0.7070.90; mean 71 SE) for the 250 m sector, from 0
times a third generation, which use alternative crucifer- to 38.78 ha (6.3475.52) for the 1250 m sectors and
ous species as hosts. The pod midge is assumed to be a 0–35.80 ha (5.6375.28) for the 1750 m sectors. Overall,
weak flyer, dispersing no more than a few hundred the area of single OSR fields ranges from 0.11 to
metres from its emergence site (Alford et al., 2003; Paul, 18.87 ha (2.2472.55). We used ARC-GIS 9.1 and
2003). ArcView GIS 3.3 (ESRI Redlands, CA, USA) for
analysing geographical data.
Species data
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 use upwind anemotaxis to locate OSR fields in spring
highest R2 for the W and NW direction for the stem (Evans & Allenwilliams, 1994, Williams et al., 2007).
weevils, whereas the pod midges revealed a weak but Prevailing winds and odour-induced upwind anemotaxis
significant reaction to the E sector. drive directional dispersal of mobile species that use
olfactory cues for resource location (Vickers, 2000),
because resource patches in upwind position have a
Discussion higher probability of being detected than resource
patches located downwind. One main process that
Our results showed that wind direction and OSR area determines the abundance of pollen beetles in OSR
specifically affected densities of the three pest species. fields is the annual dispersion from overwintering to
The response of pollen beetle density to OSR area feeding and reproduction sites. If we assume that this
showed a distinct polar pattern suggesting that beetle dispersal process is mainly guided by anemotactic search
density was predominantly affected by resource avail- behaviour, the bulk of individuals will disperse upwind
ability in the downwind landscape, whereas the upwind and touch down as soon as they encounter an OSR field
landscape seemed to be of marginal importance (Figs. 2 (Fig. 5, cf. Hein, Pfenning, Hovestadt, & Poethke, 2004).
and 4). This indicates that wind direction and resource The density of individuals at a given site will, hence,
distribution interactively affected the searching beha- depend on the OSR area along the line between the
viour of pollen beetles and confirms findings that they respective site and the overwintering habitat: the more
ARTICLE IN PRESS
D. Moser et al. / Basic and Applied Ecology 10 (2009) 208–215 213
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