International Journal of Entomology and Nematology

IJEN
Vol. 3(1), pp. 051-061, February, 2017. www.premierpublishers.org.ISSN: 2326-7262

Research Article

Behavioral and preventative management of Drosophila

suzukii Matsumura (Diptera: Drosophilidae) in Maine
wild blueberry (Vaccinium angustifolium Aiton) through
attract and kill trapping and insect exclusion-netting
Gabriel Alnajjar 1*, Judith Collins1, and Francis A. Drummond1,2
1
University of Maine, School of Biology and Ecology, Orono, ME, USA.
2
University of Maine, Cooperative Extension, Orono, ME, USA

The management of spotted wing drosophila (SWD) (Drosophila suzukii Matsumura) in small fruit
agriculture continues to receive significant attention. While complementary monitoring and
insecticidal application procedures can provide effective protection against this insect pest in
some cropping systems, more sustainable alternatives are currently under development. In this
investigation, we explored two methods of managing D. suzukii in Maine wild blueberry
(Vaccinium angustifolium Aiton): 1) the efficacy of attract and kill trapping with volatile semio-
chemicals and 2) the use of insect exclusion netting. We found that exclusion netting provided a
high degree of protection against SWD larval infestations. Consistent and significant reductions
in larval infestations were observed during field trials conducted in 2014, 2015, and 2016. The
second method, attract and kill trapping, appeared to reduce infestations in a preliminary trial in
2013. However, this result was not observed in a more thorough, replicated study in 2014. The
capture rates of traps increased with trap deployment density in blueberry plots, suggesting that
more traps per unit area concentrate flies in the local trapped area. As a result, a correlation
existed between adult captures and larval infestation despite a contact poison (boric acid) coating
the outside surface of the traps.

Keywords: Lowbush blueberry, wild blueberry, North America, Drosophila suzukii, Spotted Wing Drosophila, mass
trapping, attract and kill, insect exclusion netting

INTRODUCTION

Drosophila suzukii Matsumura, commonly referred to as contamination in North America have been estimated in
the spotted wing drosophila (SWD), is a recently various Western Pacific U.S. regions, and amounted to
established invasive insect pest of continental North roughly $US 42.9 million in three California counties
American and European small fruit cultivation. The main alone (Bolda et al., 2011).
damage caused by this pest is due to feeding of larvae
inside the fruits. However, ovipositor insertions can also
provide routes for secondary microbial invasion. The
damage potential of this exotic pest in crop systems of *Corresponding author: Gabriel Alnajjar, University of
foreign regions drastically exceeds the criterion for quality Maine, School of Biology and Ecology, Orono, ME, USA.
fruit marketability. Revenue losses attributed to SWD E-mail:gabrielalnajjar@Yahoo.com
Behavioral and preventative management of Drosophila suzukii Matsumura (Diptera: Drosophilidae) In Maine wild blueberry (Vaccinium angustifolium Aiton)
through attract and kill trapping and insect exclusion-netting
Alnajjar et al. 052
Prior to the 2011 establishment of SWD in Maine,
over 24,300 ha of wild (lowbush) blueberry (Vaccinium
angustifolium Aiton) farmland generated roughly $250
million of farm gate revenue annually between 2007 and
2012 (Yarborough, 2013). Currently, no estimate of
monetary loss in wild blueberry has been developed to
include the direct SWD infestation of fruit, secondary
microbial contamination, or the compounding costs of
implementing monitoring and treatment protocols;
although, in 2012 an estimate of 25% crop loss due to
SWD is suspected (Drummond, pers. comm.). A widely
adopted response to male detection during adult
monitoring surveys entails the targeting of adults through
application of pyrethyroid, organophosphate or spinosyn
class insecticides to suppress population growth before
significant crop damage is inflicted (Bruck et al., 2011;
Collins and Drummond, 2015 a;b). Unfortunately, many
of these insecticides threaten the health of wild and
managed pollinator species, insect natural enemies, and
persist in surrounding landscapes and watersheds after
runoff or drift dispersal (Hester et al., 2001; Beketov et
al., 2013). The development of environmentally and
economically sustainable SWD integrated pest manage-
ment (IPM) programs is currently an ongoing process in
North American and European small fruit agriculture.
Ideally, findings will be translatable across a wide
spectrum of agro-ecosystems given the species
generalist phytophagy, high generational turnover and
reproduction rates that aid in its rapid spread and
alarming crop injury potential (Asplen et al., 2015).
The use of capture-and-kill traps emitting volatile
semio-chemicals constitutes an effective chemical pre-
treatment monitoring tool of early stage SWD invasions
throughout the fruit ripening and harvest periods.
Currently, exploiting the positive chemotactic response of
SWD adults to ethanol molecules emitted by yeast-
containing baits is a widely implemented bait attraction
protocol (Walsh et al., 2011; Burrack et al., 2015). The
utilization of physiologically attractive compounds in mass
capture and kill of SWD was first proposed by Kanzawa
(1934). Since that time, results produced by two recently
conducted mass-trapping efficacy evaluations have
yielded conflicting results (Wu et al., 2007; Hampton et
al., 2014). Thus, the implementation of this approach in
the absence of complimentary control measures is widely
regarded as a high-risk option for reducing the
abundance of SWD larvae in fruit.
By exploiting the phago-stimulatory reflex of
insects to desirable food (Cevik and Erden, 2012), the
infusion of boric acid with aqueous sucrose solutions has
provided an effective oral exposure mechanism for toxin
delivery in both urban and disease vector associated
pests. Insecticidal treatments in environments in and
Behavioral and preventative management of Drosophila suzukii Matsumura (Diptera: Drosophilidae) In Maine wild blueberry (Vaccinium angustifolium Aiton)
through attract and kill trapping and insect exclusion-netting
around homes have received scrutiny due to the high
degree of direct spatial overlap between humans and
target insects. Research has shown that boric acid
ingestion by the German cockroach (Blattella germanica
L.) and the Asian tiger mosquito, Aedes albopictus
Skuse, can suppress population growth through a
combination of chronic and acute toxicity. Naranjo et al.
(2013) also described a reduction in the post-treatment
oviposition rate of A. albopictus. According to Markow
and Grady (2008), reproductive maturation of Drosophila
melanogaster females progresses more rapidly when
nutritional resources are available for immature fly
consumption. Given that recent laboratory observations
on SWD ovarian development (Alnajjar, 2016) provides
evidence for a 5-6 day reproductive immaturity in general
female flies, the ingestion of boric acid by foraging SWD
females during this critical developmental stage might
physiologically disrupt or delay the progression of repro-
ductive maturity. Hampton et al. (2014) found that a
majority of flies in close proximity to trapping grids
exclusively contact the external surface of traps and
remain free to propagate. Considering this, the
incorporation of a boric acid and sucrose solution on the
exterior surface of baited traps could expose a proportion
of these non-captured SWD to the toxin and ultimately
enhance the control efficacy of attract and kill trapping. In
the absence of a gustatory rejection by SWD upon the
ingestion of toxins in a food source, this strategy might
provide a mechanism for increasing the proportion of flies
treated upon contact with a trap surface.
Another preventative approach involves the
deployment of fine mesh netting at ripening and pre-
harvest crop intervals as a physical barrier to ovipositing
SWD contacting potential host fruits. This technique has
been effectively implemented in IPM programs for a
variety of insect pests (Lloyd et al., 2005; Dib et al., 2010;
Sauphanor et al., 2012), and has received attention in the
ongoing efforts to provide organic growers with chemical
management alternatives for SWD in a variety of
agricultural systems (Cormier et al., 2015; Schattman,
2015). The work of Cormier et al. (2015) also addressed
the significance of net shading on plant photosynthetic
activity by quantifying the chlorophyll and sugar
composition of blueberry fruits upon concluding the study.
They found no apparent reduction in fruit quality from
berries sampled in protected vs. unprotected crops.
However, given the spatial limitations of the field and
greenhouse evaluations conducted thus far, it is unknown
whether exclusion netting can meet the criterion for
economic cost effectiveness of SWD management in
large-scale crop systems. Neither of the proposed
management approaches has yet been demonstrated to
provide a level of efficacy against SWD infestations

needed in large-scale crop systems.
Research objectives
We conducted two independent field studies with the
intention of further exploring the SWD control efficacy of
these techniques in Maine wild blueberry, specifically for
small-scale growers. The first study conducted in 2013
and 2014 involved quantifying the severity of fruit
infestations in response to boric acid applications on
ethanol emitting traps deployed at varying densities
within trapping grids (attract and kill tactic). The second
investigation conducted in 2014, 2015, and 2016 exam-
ined the efficacy of insect netting in wild blueberry fruits
against SWD larval infestation.
MATERIALS AND METHODS
Attract and kill trapping
Field studies were designed to test the potential of an
attract and kill trapping tactic for wild blueberry. The
studies were conducted in wild blueberry crop fields at
the University of Maine Blueberry Hill Experimental Farm
in Jonesboro, Maine USA in 2013 and 2014. The traps
utilized in both years consisted of red Solo cups with
seven, 3.2 mm-diameter holes punched about 2.5 cm
below the cups upper rim. Within experimental grids,
individual traps were positioned on a green, metallic 76
cm plant support post and baited with approximately 5 cm
of a mixture containing 15 ml yeast granules, 60 ml white
granulated sugar, and 0.35 L H2O. Control traps were
filled with about 5 cm of water only. We used sugar
syrup/yeast bait since it has proven to be a superior
attractant in wild blueberry (Burrack et al., 2015). Each
red Solo cup was covered with a red foam photo-
absorption cap to seal the trap and prevent the
interference of light with fermentation reactions. The
external surface of each experimental trap was then
uniformly sprayed with a mixture of 1% borate L -1 25%
(w/v) sucrose/water solution. Traps intended for
monitoring purposes during these assessments did not
have boric acid solution sprayed on the external surfaces.
Fruit samples were collected to assess larval infestation.
Each fruit sample occupied a volume of about 236 ml,
and was gathered from random wild blueberry clones
(genets) within each treatment plot. All fruit samples were
processed within one week after collection. Each sample
was gently crushed in a plastic bag and suspended in
10% saline solution for 30 min in order to induce
disassociation of SWD larvae from fruit pulp. The
contents were then filtered through a fine mesh sieve and
Behavioral and preventative management of Drosophila suzukii Matsumura (Diptera: Drosophilidae) In Maine wild blueberry (Vaccinium angustifolium Aiton)
through attract and kill trapping and insect exclusion-netting
Int. J. Entomol. Nematol. 053
suspended in a black bottom metallic tray filled with
water. A lamp was utilized to illuminate the contents of
samples so the abundance of SWD larvae inhabiting fruit
samples could be determined (Drummond et al. 2016).
Red cup traps were gathered at different intervals
depending on the experiment, and analyzed within one
week of being collected. The contents of each trap were
rinsed with H 2O and filtered through a sieve in the
laboratory. The remaining trap contents were emptied
into a white metal tray filled halfway with about 5 cm
water. Male and female SWD counts were then obtained
from each trap sample.
On 25 July 2013, a single 7.6 m x 4.9 m study
area was set up at the University of Maine Blueberry Hill
Experiment Station in Jonesboro, ME, for a preliminary
investigation of the potential of mass trapping for
suppression of SWD in pre-harvest fruits. Twelve traps
were deployed in a 3 x 4 grid with approximately 1.8 m of
spacing between traps. Traps were collected and
replaced with freshly baited traps every seven days.
Those containing flies were taken back to the laboratory
where the abundance of male and female SWD was
determined.
On the final collection date, 14 September 2013,
two samples of blueberries were gathered from random
areas inside and outside the grid to determine female
SWD egg laying activity. Externally sampled plant clones
were located a minimum of 9.1 m from the peripheral
border of the trapping grid. Each blueberry sample was
processed in the laboratory by using the methods
described previously.
A replicated field study was completed in 2014.
On 27 August 2014, prior to the detection of juvenile
SWD in preliminary fruit monitoring bouts, twelve, 9.1 m x
9.1 m study grids were established in wild blueberry.
Experimental traps were deployed and spaced at 0.9 m
(low density), 1.8 m (medium density) or 2.7 m (high
density) in order to determine the impact of varying trap
density on SWD abundance. Control traps were spaced
1.8 m apart. All treatment grids were replicated three
times, in replicate blocks in the same 16.2 ha field at the
University of Maine Blueberry Hill Experiment Station in
Jonesboro, ME. Trapping grids within a block were
positioned a minimum distance of 9 m from one another.
For collection of adult abundance data, three randomly
chosen monitoring traps were taken from each trapping
grid. During weekly trap collections, all traps were
cleaned and recharged with freshly prepared bait and
boric acid solution. In addition, three blueberry samples
(236 mL each) of random clones located within each
study area were collected weekly so that larval
infestations could be quantified.
Since treatments were not replicated in the
preliminary attract and kill trapping study, assessment in
Alnajjar et al. 054
2013, only the 2014 study results were statistically
analyzed. Repeated measures analyses of variance
(ANOVA, RCB, repeated measures are 4 sampling times)
were utilized to examine the impact of treatment and
block on the observed variation in adult and larval SWD
abundance within trapping grids. Analysis was conducted
on square root transformed dependent variables.
Analyses of variance (ANOVA, RCB) were utilized to
examine the impact of treatment and block on the
observed variation in adult and larval SWD abundance
within trapping grids. An initial test entailed two
separately conducted ANOVAs to assess how varying
trap density impacted SWD adult and larval abundances
within study plots. A third ANOVA examined the relative
abundance of SWD larvae found infesting fruit samples
collected in control compared to medium density
experimental plots. Mean separation was performed
using a Turkeys post-hoc test (JMP , 2015).
Insect exclusion netting
Studies conducted in 2014, 2015, and 2016 were de-
signed to test the effectiveness of minimizing SWD larval
infestation of fruit by preventing female SWD adults
access by covering the crop with netting. Anti-Insect
Netting (#25 Mesh) with mesh size meeting the pre-
determined dimensions for effective exclusion of SWD
adults (Caprile et al., 2013) was tested at Blueberry Hill
Farm Experiment Station in Jonesboro, ME during the
summers 2014 and 2015. In 2016, a finer, but less
expensive mesh netting, Agribon + AG-19 Row Cover
(commonly referred to as Reemay, a spun polyester
material) was used to protect fruit from SWD attack.
In 2014 there were two replicated netting en-
closures (each 3.96 m x 15.24 m), one replication was
initiated on 9 July and a second on 18 July; in 2015, two
netting enclosures (each 3.96 m x 15.24 m) were set up
on 29 July, with a third (3.96 m x 30.48 m) initiated on 30
July. In 2016, two netting enclosures (each covering an
area of 15.25 m x 45.7 m) were set up on 14 July. All of
these replicates were deployed prior to capture of any
adult SWD and prior to 0-5% of fruit being susceptible to
attack (ripe fruit). An adjacent non-protected plot served
as a control treatment for each replicate. In addition,
ethanol baited, red cup, monitoring traps were deployed
in close proximity to study areas to determine the onset
of invasions. Upon initial capture of adult SWD,
blueberries were periodically sampled from the non-
protected study plots in order to track fruit infestation.
After fruit was naturally infested by SWD, exclusion nets
were removed and fruit samples were collected to
determine the abundance of SWD larvae. In 2014 and
2015, five fruit samples were taken from areas protected
by the netting and paired areas not protected by any
Behavioral and preventative management of Drosophila suzukii Matsumura (Diptera: Drosophilidae) In Maine wild blueberry (Vaccinium angustifolium Aiton)
through attract and kill trapping and insect exclusion-netting
netting (sample size was 472 and 157 ml in 2014 and
2015, respectively). In 2016, 10 samples (236 ml each)
were obtained from each study plot. Samples were
assessed utilizing the previously described salt extraction
method for fruit samples. In addition, in 2016 fruit
ripeness and yield were qualitatively assessed between
the non-netted control and the net enclosure treatments.
A randomized block design model, with year as a
blocking factor, was utilized for analyzing the larval fruit
infestation (adjusted to larvae per 236 ml of fruit) over the
three years (JMP , 2015).
RESULTS
Attract and kill trapping
In 2013, there was a considerable increase in the
abundance of SWD flies captured in baited traps toward
the end of the preliminary mass trapping experiment (Fig.
1), with 373.4 188.2 (SD) flies/ trap on 14 September
2013. The majority of SWD flies were females. Fruit
collected within the trapping grid contained 68.3 SWD
larvae per sample in comparison to 125.4 larvae found
infesting fruits obtained from the paired plot that was not
trapped (Fig. 2). This corresponds to a 46% reduction in
larval abundance, with a negligible 0.8 g difference in fruit
sample weights of berries obtained from blueberry clones
in trapped compared to non-trapped treatments. Despite
the lack of replication, this preliminary experiment
suggested that a high density of traps has the potential to
reduce the infestation of berries by SWD.
In the 2014 replicated trapping out study,
comparing control grids (1.8 m trap spacing) with the
baited trapping grids of medium density spatial
arrangement (1.8 m trap spacing), there was no
statistically significant difference in larval abundance over
time in blueberry samples (F (1,19) = 0.12, P = 0.97). No
adult SWD were captured in the non-baited control traps
compared to 17.2 21.1 (SD) SWD per trap captured in
the medium density treatment. Figure 3 shows that
varying trap spacing of experimental grids did have a
significant effect on the larval infestation in blueberry
samples (F (1,30) = 15.00,P = 0.001)..
Larval infestation increased with increasing trapping
intensity suggesting that female SWD flies were attracted
into the plots at a higher frequency. While there was no
significant difference detected in the mean abundance of
adults captured in traps across the range of trap-densities
(F(1,30) = 0.74, P = 0.40), a trend of increasing fly trap
captures with increasing trapping intensity can be seen in
Figure 3. This trend in adults captures with trapping
intensity is correlated with the abundance of larvae in the
trapped plots (r = +0.631,P < 0.0001).
 Int. J. Entomol. Nematol. 055

Figure 1. Abundance of SWD flies captured in each of twelve traps constituting a single
trapping grid located in Jonesboro, ME during the summer of 2013.

Figure 2. Abundance of SWD larvae infesting fruit samples gathered within a trapping grid plot
or in a control plot, experimental site located in Jonesboro, ME during the summer of 2013.

Insect exclusion netting ml) from non-protected blueberry plots among the 2014,
2015, and 2016 trials (P = 0.734), nor was there any
There was no difference in mean larval infestation of 9.7 difference between mean counts of 0.29 0.29, 0.13
7.7 (se), 2.2 0.4, and 5.9 2.1 per fruit sample (236 0.07, 0.20 0.10 larvae obtained from net protected plots
Behavioral and preventative management of Drosophila suzukii Matsumura (Diptera: Drosophilidae) In Maine wild blueberry (Vaccinium angustifolium Aiton)
through attract and kill trapping and insect exclusion-netting
Alnajjar et al. 056

Figure 3. Mean abundance of individual D. Suzukii captured in control, low density (2.7 m),
medium density (1.8 m) and high density (0.9 m) experiment trapping grids. Each mean represents
the average of three replicates over four weeks of trapping. Letters above larval abundance
columns represent Tukey post-hoc results, with bars displaying dissimilar letters signifying
statistical significance.

Figure 4. Mean abundance of D. suzukii larvae inhabiting five blueberry samples (adjusted to 236 ml each) from uncovered control plots or crop plots
protected with exclusion netting. The measurements presented here represent the combined data from three consecutive trials conducte d in 2014,
2015, and 2016. Error bars represent the standard error of the mean

Behavioral and preventative management of Drosophila suzukii Matsumura (Diptera: Drosophilidae) In Maine wild blueberry (Vaccinium angustifolium Aiton)
through attract and kill trapping and insect exclusion-netting

Figure 5. Photograph of 2016 netting deployment and delayed fruit maturation exhibited in plots covered with Agribon row
cover (Reemay).
between 2014, 2015, and 2016; respectively (P = 0.891).
Over all three years the exclusion netting provided a high
degree of protection from larval infestation in comparison
to non-netted plots (F(1,8) = 19.958, P = 0002, Fig. 4).
In 2016 we also made qualitative observations
the quantity and quality of fruit under the Agribon row
cover (Reemay). Although this less expensive cover
proved highly effective against SWD fruit infestation, it
was difficult to deploy as the cloth fabric was easily torn.
In addition, fruit appeared to have been knocked off
stems possibly due to excessive movement of the fabric
in high winds, and some fruit was delayed in maturation
(red not blue), but there was no evidence of diseased fruit
(Fig. 5).
DISCUSSION
After experimentation with attract and kill trapping and
insect netting in wild blueberry, only preventative
exclusion of adult SWD with netting demonstrated
effective management potential. Over all three years,
netting greatly suppressed infestation of wild blueberry
fruit. These results complement the findings of Cormier et
al. (2015) who reported zero SWD adult emergences
from high bush blueberry fruits grown under net-protected
plots. Furthermore, this approach is thought to entail a
high degree of SWD exclusion efficacy in other cultivated,
small-fruit producing plants such as raspberry
(Schattman, 2015). To date, all positive results with this
technique have been obtained from studies conducted
under limited space where the area of the study plots
represents only a fraction of crop coverage necessary in
large-scale production operations. Increasing the land
Behavioral and preventative management of Drosophila suzukii Matsumura (Diptera: Drosophilidae) In Maine wild blueberry (Vaccinium angustifolium Aiton)
through attract and kill trapping and insect exclusion-netting
Int. J. Entomol. Nematol. 057
area for protection would substantially increase the
amount of netting product, labor and maintenance inputs
required.
Excluding labor requirements for seasonal
installation and maintenance of netting, the deployment
of the most inexpensive exclusion netting (Agribon ) on a
single hectare would cost roughly US $4,600 according to
product pricing by Gardeners Supply Company.
Therefore, relative to Anti-insect Netting at US $6.09 /
2
13 ft or US $49,680 per hectare, Agribon appears to be
a cost effective alternative. According to Chen et al.
(2015), from 2010 to 2014 organic wild blueberry
cultivation in Maine generated an annual net revenue of
US $3,724 per hectare. In a single year, therefore, the
financial inputs of purchasing netting units alone would
exceed the total amount of generated revenue if the
Agribon netting was used. However, more economically
justifiable cost estimates are derived when considering
the repeated use of netting. Assuming a degree of
material durability and longevity spanning 2, 3, 4 and 5
years, the respective proportional cost decreases to
roughly 62%, 41%, 31%, and 25% of total revenue
generated from organically produced wild blueberry.
However, our experience with the Agribon material
suggests only a one year use for a two month duration
during the growing season. Given the high degree of
infestation mitigation obtained with exclusion netting
reported in this and other studies, it might be more
feasible to deploy this management tactic if the yield
damage caused by SWD exceeds the depreciated annual
cost estimate of using a single netting unit repeatedly
over time. Presumably, this disparity will increase over
longer re-use time periods. Unfortunately, monetary
losses due to SWD infestation have not yet been
Alnajjar et al. 058
quantified in the wild blueberry crop system; although,
some farms have experienced up to 25% crop loss due to
SWD in some years (Drummond, pers. comm.).
Therefore, this technique has great promise only if
multiple-year use of netting is possible.
Insect exclusion netting has also demonstrated
considerable success against plant pathogen vectors,
preventing their physical contact with host plants in order
to suppress insect-mediated disease transmission.
Stanley et al. (2004) demonstrated the potential for using
this technique to limit the spread of tomato yellow leafcurl
virus (TYLCV) among greenhouse tomato plants. Plants
displayed TYLCV symptoms less frequently when
protected with netting for exclusion of the pathogens
insect vector, the whitefly Bemisia tabaci (Gennadius).
Another relevant phenomenon has been described by
Louise et al. (1996) for Drosophila melanogaster and the
dispersal of its microbial symbiont Botrytis cinerea, a
necrotrophic fungus responsible for the occurrence of
grape rot in vineyards. External localization of fungal
spores on fly integuments, in conjunction with internal
occupancy of the flies alimentary canal is thought to
facilitate passive spore dispersal throughout vineyards
harboring D. melanogaster populations (Louise et al.,
1996).
The symbiotic association of SWD with Botrytis
cinerea or alternative pestiferous microbes has not been
documented, nor have the risks of introducing microbial
symbionts of SWD into recently colonized agro-
ecosystems been addressed in the literature. However,
considering the described mechanisms of passive grape
rot transmission by Drosophila melanogaster, it is
possible that symbiotic associations within SWD may
diversify the pest pressures imposed on certain plant
species that further reduce fruit marketability, perhaps
even leading to the inadvertent introduction of additional
pestiferous microbes into an agro-ecosystem.
The exclusion efficacy of netting in preventing
SWD flies from contacting viable fruit hosts has been
demonstrated in multiple crop systems. Coupled with the
theoretical cost-efficiency in the context of Maine wild
blueberry production, it is justifiable to consider exclusion
netting as a viable avenue for further evaluation as a
component of developing IPM programs for SWD.
However, one potential limitation to this method of SWD
prevention includes the alteration of microclimates
located within netting units. If the accumulation and
retention of moisture within the space encapsulated by
the netting material effectively alters crop and pest
growth conditions in relation to ambient environmental
conditions, the consequent humidification could create a
habitat in which plant pathogenic fungi flourish (Ciancio
and Mukerji, 2008). Although not yet tested for, this is a
necessary consideration to address should the technique
Behavioral and preventative management of Drosophila suzukii Matsumura (Diptera: Drosophilidae) In Maine wild blueberry (Vaccinium angustifolium Aiton)
through attract and kill trapping and insect exclusion-netting
be considered for SWD management in large-scale crop
systems where fungal pests may become problematic
and require damage control. We have not observed any
increased fungal disease in wild blueberry due to the use
of netting, but particularly wet summers could result in an
increase in disease incidence.
We also evaluated attract and kill trapping as a
tactic for reducing the quantity of SWD larvae infesting
berries. In comparison to control study grids with traps
containing water, deployment of traps containing an
ethanol-emitting bait and topical boric acid application
failed to reduce the abundance of larvae infesting
blueberry samples. Our study provided evidence that
increasing trap density in wild blueberry plots attracted a
concomitant increasing number of flies to a study area.
This aggregation effect, whereby the number of flies in
the proximity of trapping grids increases proportionally
with trap density, would have consequently increased the
frequency of contact between egg laying females and
healthy blueberry fruits. Alnajjar (2016) found that these
same traps baited with sugar syrup and yeast only catch
approximately 16% of flies restricted to a localized area
of wild blueberry with field cages. The capture efficacy of
a comparable trap design was assessed by Hampton et
al. (2014) who found variable capture rates ranging from
10% - 30% in traps baited with a combination of acetic
acid and ethanol. Although the impact of boric acid was
not quantified in this study and may have negated the
fitness of some non-captured individuals, it is also
possible that the presence of boric acid acted as a
repellent. Regardless of this, our attract and kill study
did not achieve satisfactory control. Hampton et al.
(2014) suggested the incorporation of insecticides directly
on the external surface of traps, and/or on proximal
plants as exposure reservoirs. Doing so might eliminate
the attracted, non-captured adults given the high efficacy
of various insecticides against SWD (Bruck et al., 2011;
Collins and Drummond, 2016a; 2016b). The results of
Hampton et al. (2014) also imply an inverse relationship
between distance from trap and the probability of SWD
infestation. It has therefore been proposed that trapping
grids be deployed on the periphery of managed fields in
order to confine invasions within trap cropping reservoirs
in these areas so they can be effectively treated and
prevented from further dispersal into the fields during pre-
harvest intervals.
Additional evaluations on SWD bait preference
and the chromatic attractiveness of traps have provided
insights for the development of trap designs that enhance
SWD fly attraction and capture rates. The physical
localization of SWD flies on the exterior portion of traps is
believed to occur more frequently on dark pigmented
surfaces. Basoalto et al. (2013) reported double the
observed fly capture rates in traps wielding a black

painted strip enveloping entry holes for the flies. The
implementation of this trap coloration protocol in SWD
monitoring may lead to earlier and more accurate
detection of flies in agricultural landscapes (Basoalto et
al., 2013) and also enhanced proficiency in the capture
rates of individual traps within mass-trapping grids. Given
the innate implications of nutritional, copulatory and
reproductive rewards the overall attractant power of traps
is mainly influenced through primary physiological
attraction of drosophilid flies to naturally occurring volatile
semio-chemicals, (Landolt et al., 2011). Potentially then,
developing a trap that is visually appealing to SWD flies,
combined with baiting with optimal chemical mixtures for
spatially expansive and powerful attraction, could
constitute a more effective mass-trapping management
tactic with greater SWD control potential than the
protocols tested here.
Field trials on the preference of SWD with various
volatile compounds by Landolt et al. (2011, 2012)
suggest that the maximum positive chemotactic response
of adult SWD occurs during concurrent emission of
ethanol and acetic acid, as indicated by a greater
abundance of adults captured in traps emitting both baits
in comparison to those containing either compound
independently. However, the SWD chemosensory niche
specialization for reproductive exploitation of fresh or
ripening fruits in nature appears to be in stark contrast to
the mechanics of physiological stimulation in Drosophila
melanogaster. This latter species displays a consistent
positive chemotactic response upon detection of volatile
fermenting byproducts emitted by decaying plant
material. Keesey et al. (2015) conducted a laboratory
bioassay in which olfactory attraction cues for SWD and
its close relative, Drosophila biarmipes Malloch, were
found to positively correlate with the volatile isoprenoid -
cyclocitral. The compound -cyclocitralis is believed to
have developed a novel association with SWD neuronal
receptors. This volatile semio-chemical is associated with
the host plant leaf tissue and is therefore suspected of
significantly influencing the degree of SWD attraction
toward ripening fruits located on the stems of otherwise
healthy plants. Taken together, evaluating the exploit-
ation of -cyclocitral and other as of yet unknown
phytochemicals in attract and kill trapping of SWD should
consider the preparation of chromatically appealing trap
exteriors for increasing the capture frequency. It should
also consider the arrangement of traps at relatively low
concentrations for decreasing the probability of fly
aggregation and maximizing the degree of management
capability that can be achieved with this technique. For
the purposes of trap cropping, however, it may be the
case that maximizing the concentration of flies in trap
crops is more desirable given that a greater quantity of
flies can be targeted with a single insecticide treatment.
Behavioral and preventative management of Drosophila suzukii Matsumura (Diptera: Drosophilidae) In Maine wild blueberry (Vaccinium angustifolium Aiton)
through attract and kill trapping and insect exclusion-netting
Int. J. Entomol. Nematol. 059
ACKNOWLEDGEMENTS
We would like to thank Dr. Andrei Alyokhin and Dr.
Eleanor Groden for reviewing a previous version of this
manuscript and Ms. Tamara Levitsky, Ms. Elissa
Ballman, Ms. Jen Lund, Mr. Michael Hahn and Mr.
Andrew Wilson for their assistance with the research
conducted in the laboratory and field. We also would like
to acknowledge Mr. Jeff Brann, Mr. Josh Stubbs, and Mr.
Chris McMannus at the University of Maine Blueberry Hill
Research Farm in Jonesboro, ME for their help with this
research. This is Maine Agriculture and Forestry
Experiment Journal number 3508. Partial funding for this
project was provided by the National Institute of Food and
Agriculture, U.S. Department of Agriculture Specialty
Crops Research Initiative under Agreement No. 2015-
51181-24252.
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Accepted 12 November, 2016

Citation: Alnajjar G, Collins J, Drummond FA (2017).
Behavioral and preventative management of Drosophila
suzukii Matsumura (Diptera: Drosophilidae) in Maine wild
blueberry (Vaccinium angustifolium Aiton) through attract
and kill trapping and Insect exclusion-netting, Inter-
national Journal of Entomology and Nematology 3(1):
051-061.

Copyright: 2017 Alnajjar et al. This is an open-access

article distributed under the terms of the Creative
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use, distribution, and reproduction in any medium,
provided the original author and source are cited.

Behavioral and preventative management of Drosophila suzukii Matsumura (Diptera: Drosophilidae) In Maine wild blueberry (Vaccinium angustifolium Aiton)
through attract and kill trapping and insect exclusion-netting