ASPEN-00295; No.

of pages: 5; 4C:
Journal of Asia-Pacific Entomology xxx (2011) xxx–xxx

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Journal of Asia-Pacific Entomology

journal homepage: www.elsevier.com/locate/jape

Distribution, spread, and impact of the invasive hornet Vespa velutina in South Korea
Moon Bo Choi a, Stephen J. Martin b, Jong Wook Lee a,⁎
a
Department of Life Sciences, Yeungnam University, Republic of Korea
b
Department of Animal & Plant Sciences, University of Sheffield, Sheffield, S10 2TN, UK

a r t i c l e i n f o a b s t r a c t

Article history: Hornets (Vespa spp) are top insect predators that can control pests, but their venomous stings and defensive be-
Received 21 September 2011 havior cause numerous human deaths throughout Asia. Hornets usually inhabit rural areas which reduces poten-
Revised 24 November 2011 tial conflict with humans. In 2003, the invasive hornet, Vespa velutina, arrived in southern Korea (Yeongdo
Accepted 26 November 2011
region) and became established. It is currently spreading northwards at a rate of 10–20 km per year. Despite
Available online xxxx
originating in tropical/subtropical areas of Indo-China, its nesting biology and life cycle in South Korea are similar
Keywords:
to those found throughout its native range, with mature colonies containing 1000–1200 adults. In 7 years, V.
V. velutina velutina has become the most abundant hornet species in Southern Korea by displacing native Vespa species
Vespa such as V. simillima, which has a similar nesting biology. We also found a significant positive correlation between
Invasive the abundance of V. velutina and the degree of urbanization, indicating that this invasive species was well
Spread adapted to urban environments. This was supported by our finding that 41% of emergency call-outs (119 Rescue
Urbanized Services) to deal with social wasps/hornet problems were due to V. velutina, which was twice as high as the num-
Busan ber of calls about the next most abundant species. The rapid spread of V. velutina across southern Korea indicates
that this species will continue to spread north-westward in the Korean peninsula and will become a major prob-
lem as more people and beekeepers come into contact with this aggressive invasive hornet.
© Korean Society of Applied Entomology, Taiwan Entomological Society and Malaysian Plant Protection Society,
2011. Published by Elsevier B.V. All rights reserved.

Introduction
Climate change is predicted to alter the range of many species, in-
cluding allowing the expansion of invasive species (Hill et al., 2010). A
tropical/subtropical species of hornet, V. velutina nigrithorax, was
recorded for the first time in South Korea in 2003 (Kim et al., 2006)
and in France in 2004 (Haxaire et al., 2006). Both of these countries
have a temperate climate. Their subsequent rapid spread has impact-
ed human activities and the native fauna. V. velutina is now considered
a serious invasive pest in France (Chauzat and Martin, 2009; Haro et
al., 2010) and potentially in South Korea. The native range of V. velu-
tina is the tropical and subtropical regions throughout most of Indo-
China, Indonesia, and Taiwan (Archer, 1994; Martin, 1995; Carpenter
and Kojima, 1997). V. velutina normally occurs in the cooler mountain-
ous highland areas throughout its native range (Martin, 1995) and so
may be pre-adapted to exploit temperate environments. The southern
parts of South Korea are undergoing environmental (e.g. Kim, 2007)
and vegetation (Park et al., 2010) changes, resulting in a more sub-
tropical climate. Correspondingly, ten different subtropical butterfly
species are now recorded in South Korea (Park et al., 2006) and the
tropical cricket, Lycorma delicatula, has spread throughout the country
since 2004 (Korea Forest Research Institute, 2007).
V. velutina adults can catch honeybees in flight as they return to the
colony. The native Asian honeybees (Apis cerana) have evolved a ‘heat-
balling defense’ warning behavior when hornets are patrolling near
their nest entrance (Tan et al., 2005). However, European honeybees
(Apis mellifera), which are commonly reared by Korean beekeepers,
do not possess this behavior and are more susceptible to attack from
V. velutina (Shah and Shah, 1991; Abrol, 1994; Tan et al., 2007). South
Korea has six native hornet species, V. analis, V. mandarinia, V. simillima,
V. crabro, V. ducalis, and V. dybowskii. The adult size, nest structure, nest-
ing habits, and population size of V. velutina are very similar to V. simil-
lima, a species that inhabits more temperate regions of Asia, including
South Korea (Matsuura, 1973; Martin, 1995). Therefore, V. simillima
may be in direct competition with V. velutina. Even in its native range,
V. velutina is one of the most aggressive and feared hornets (Martin,
1995) and may pose a serious risk to humans if they move into more
urban areas. The purpose of this study is to examine the spread of V.
velutina across South Korea during the past 7 years, and its habitat pref-
erences, damage, and impact on native Vespa composition.
Materials and methods
Study area

In 1997 to 2001, all native hornet colonies or adults were surveyed

⁎ Corresponding author. by trapping and netting in the Busan city area. After the arrival of V.
E-mail address: jwlee1@ynu.ac.kr (J.W. Lee). velutina in 2003, nine cities (Yangsan, Ulsan, Jinhae, Masan, Gyeongju,

1226-8615/$ – see front matter © Korean Society of Applied Entomology, Taiwan Entomological Society and Malaysian Plant Protection Society, 2011. Published by Elsevier B.V. All
rights reserved.

Please cite this article as: Choi, M.B., et al., Distribution, spread, and impact of the invasive hornet Vespa velutina in South Korea, J. Asia Pac.
Entomol. (2011), doi:10.1016/j.aspen.2011.11.004
2 M.B. Choi et al. / Journal of Asia-Pacific Entomology xxx (2011) xxx–xxx

Cheongdo-gun, Haman-gun, Gyeongsan and Daegu) within 200 km of

Busan city were monitored by trapping adult hornets. During 2010,
we conducted a detailed study in Busan city by trapping and netting
adults at 75 locations throughout Busan city that represented all
grades of urbanization. We also collected adults of all destroyed hor-
net nests collected by 119 Rescue Services (emergency call-outs)
within the Geumjeong-gu region of Busan city. Busan city has a tem-
perate climate that is strongly influenced by oceanic currents. It is lo-
cated at the far southeastern end of the South Korean peninsula. It is a
large city with a population of 3.5 million, and is home to the coun-
try's largest trading port. It is surrounded by mountains and forests
and has many parks and green areas that provide excellent habitats
for many species of hornets.

Classified urbanized grade

The distribution of hornets is greatly influenced by the availability of

nesting sites, which fall into two main categories: (i) underground or
enclosed (cavity) nests representative of V. mandarinia and V. crabro,
and (ii) open nests typical of V. analis, V. simillima, and V. velutina
(Matsuura, 1984; Martin, 1995). Because Busan city covers a region in
which both forest and urban areas coexist, we classified the degree
urbanization into five grades for each collection area. Grading was esti-
mated by the percentage of green or urban coverage (modified percent-
age of vegetation and pavement/building cover [Zanette et al., 2005]) of
a 100 × 100 m 2 area around the 75 collection sites. Grade 1 indicates na-
ture forest regions with no artificial features; Grade 2 is the forest edges
adjacent to city with b25% urbanization; Grade 3 is the urban areas with
large wooded parks within the city or university campuses (urbaniza-
tion 50%); Grade 4 is the regions such as small community parks, river
terraces, or gardens with b75% urbanization; Grade 5 indicates down-
town urban areas where no green areas exist.

Sampling of Vespa species
In this study, Vespa species were sampled by a combination of net-
ting flying adults, collecting nests, and using traps. The trap consisted of
a 2 L plastic container containing a solution of 1:1:1 sugared water,
acetic acid, and ethanol that is attractive to hornets and widely used
by beekeepers to control hornets. The traps were hung throughout
the Busan city area and monitored every 2 weeks. During 1997–2001,
sampling was conducted via 50–60 traps and netting in Busan between
July and September of each year. During 2003–2009, the spread of V.
velutina was monitored in Busan city by installing 20 traps in all
major habitats. This was supplemented by 1–2 days of trapping and
netting in other regions outside Busan. In 2010, one trap was installed
at each of 75 locations, i.e. 15 locations for each Grade of urbanization.
In addition, three traps each were installed in the urban and forest
areas in Haman-gun, Gyeongsan city, Masan city, and Yeongyang-
gun. The Geumjeong-gu region of Busan city 119 Rescue Services
emergency call-out reports dealing with social wasp/hornet nests
problems during August–September 2010 were also studied to investi-
gate the scale of the problem.
The 1997–2001 samples were deposited in the Forensic & Conser-
vation Biology Lab at the Department of Biological Science, Kosin Uni-
versity, and all other samples are at the Animal Systematics Lab at the
Department of Life Sciences, Yeungnam University, South Korea. Sta-
tistical analyses were analyzed with correlation analysis by Pearson's
Correlation Coefficient at p b .005 using software SPSS V.16.
Results and discussion
Distribution and spread of Vespa velutina in South Korea
Since the first report in 2003, V. velutina has spread north and west
from the point of origin at a rate of 10–20 km per year (Fig. 1), as
Fig. 1. Spread and distribution of V. velutina between 2003 and 2010 both within the
Busan metropolitan city (insert) and across southern regions of South Korea. Key: a,
Busan metropolitan city; b, Yangsan city; c, Ulsan city; d, Jinhae city; e, Masan city ; f,
Gyeongju city ; g, Cheongdo-gun ; h, Haman-gun ; i, Gyeongsan city j, Daegu city; k,
Yeongdo-gu (Mt. Bongraesan); l, Seo-gu (Mt. Seunghaksan); m, Busanjin-gu
(Mt. Baekyangsan); n, Haeundae-gu (Mt. Jangsan) o, Geumjeong-gu (Mt. Geumjeong-
san); p, Gijang-gun ; q, Gangseo-gu; r, Yeongyang-gun (points on map are sampling
sites).
indicated by the arrival in 2010 of V. velutina in Daegu city, 110 km
from Busan city. It has been speculated that this species has spread
via the movement of goods before colonies first appeared around
the Busan port area (Fig. 1), the largest trading port in South Korea.
Further weight is given to this idea because the arrival of V. velutina
in France is believed to be via imported pots from China (Chauzat
and Martin, 2009). Furthermore, the same distinctive subspecies,
V. velutina nigrithorax, that occurs in Indo-China has invaded both
France and South Korea suggesting a similar point of origin.
Biology of V. velutina nigrithorax
In South Korea, V. velutina queens were observed each year
searching for sap and nest material during May (Fig. 2A), and mating
(Fig. 2D) was observed during November, indicating a 7 month nest-
ing cycle that is comparable to the native hornets V. simillima and V.
mandarinia. Like V. simillima the embryo colony of V. velutina is built
in an enclosed space and the entire colony later relocates to
an open nest-site where the colony is free to expand rapidly. During
August–September, V. velutina colonies in South Korea were
40–50 cm in height and 25–30 cm in length, with 4–5 combs. Mature
colonies located in November were 60–90 cm in height and 40–70 cm
in length, with 6–7 combs and containing 1000–1200 adult workers,
which makes them the most populous species of hornet now in

Please cite this article as: Choi, M.B., et al., Distribution, spread, and impact of the invasive hornet Vespa velutina in South Korea, J. Asia Pac.
Entomol. (2011), doi:10.1016/j.aspen.2011.11.004
 M.B. Choi et al. / Journal of Asia-Pacific Entomology xxx (2011) xxx–xxx 3

nest site preferences of different hornets. Seven years after the arrival
of V. velutina it has become established in all grades of habitat but
shows a significant positive correlation (Pearson r = .97, p b .005,
n = 5) with the level of urbanization (Fig. 3). This has resulted in V.
velutina becoming the most abundant hornet in urban areas by dis-
placing V. simillima. This is due possibly to the similarity in nesting cy-
cles and competition for nesting sites and food between V. velutina
and V. simillima. The annual temperature of urbanized areas is
2–3 °C higher than surrounding areas (Kim and Baik, 2002) due to
the heat island effect (Landsberg, 1981). However, this may not be
as important for hornets as for other insect species because hornets
are excellent at maintaining elevated internal nest temperature
relative to the ambient temperature (Martin, 1990).

Potential conflict of V. velutina with humans

V. velutina is well adapted to urban environments and is predicted

Fig. 2. Biology of V. velutina. A. Worker gathering nest materials, Pittosporum tobira,
B. Mature nest in Osmanthus heterophyllus tree. C. Destroying a colony in a downtown
urban area. D. Mating behavior in November.
South Korea. Within Busan city, V. velutina nests were found mainly in
open spaces that contained bushes and trees (e.g. Osmanthus hetero-
phyllus, Pittosporum tobira) or under building eaves (Fig. 2B, C) since
there is limited wooded area in the city. These features are all similar
to those found in the native range of V. velutina (Matsuura, 1973;
Starr, 1992; Martin, 1995).
Changes in the native hornet population associated with the establish-
ment of V. velutina
Prior to the arrival of V. velutina, the relative abundance of the six
native hornet species between 1997 and 2001 in Busan city were V.
analis > V. mandarinia > V. simillima > V. crabro > V. ducalis > V. dybow-
skii (Table 1). However, in 2010, only 7 years after the establishment
of V. velutina, it has become the most abundant species in Busan. It ac-
counts for 37% of the entire hornet population and has caused a 20%
drop in V. simillima population and a 10% drop in V. mandarinia
populations (Table 1). In other regions in southern Korea, V. crabro
is the dominant species (Table 1). However, after only 1 year in
Gyeongsan city, V. velutina has become established and represents
17% of the total hornet population. Further monitoring is required
to determine if V. velutina will displace other native hornet species
in these areas.
The level of urbanization has a strong influence on the relative
abundance of native hornet species (Fig. 3). We found V. analis, V.
mandarinia, and V. crabro dominant in forested areas (i.e. Grade 1
and 2 regions) and V. analis and V. simillima dominant in parks and
more urban areas (i.e. Grades 3, 4 and 5) (Fig. 3), which supports
the findings of Choi and Moon (2005). This pattern is largely due to
to come into conflict with humans. During August–September 2010,
there were 78 colonies of social wasps and hornets removed by the
119 Rescue Services emergency call-outs due to concerns over public
welfare in the Geumjeong-gu region of Busan city (Table 2).
V. velutina accounted for 41% of these call-outs, representing twice
as many as the next species Polistes rothneyi (Table 2), indicating
that its preference for urban environment is bringing it into conflict
with humans. Between 2000 and 2003 (prior to the arrival of V.
velutina) approximately 65–70 call-outs per year were made in the
Geumjeong-gu region of Busan city, with V. simillima and P. rothneyi
accounting for 85% of the cases (Choi and Moon, 2003). Furthermore,
Jung et al. (2008) reported that in an apiary in Busan approximately
50 out of 300 bee colonies perished due to attack by V. velutina during
a 2–3 week period, which mirrors the problems that beekeepers are
facing in France. Currently, the only control measures are removing
the colony using fire (Fig. 2C) or insecticides (Makino et al., 1981).
However, these are only a temporary means in urban areas and
they do not provide a fundamental solution to controlling the wasp/
hornet problem. Social insects perform many activities, including in-
sect pest control, that benefit mankind. Therefore, they also need to
be preserved in some way (Moon, 2001). However, this is made diffi-
cult when a dangerous invasive species settles in urban areas and
causes major problems for beekeepers.
Conclusion
Since V. velutina was first found in Busan city in South Korea in
2003, it has become well established in southern Korea and is spread-
ing at a rate of 10–20 km per year. The original arrival is believed to
be via imported goods from China, which may be how it was intro-
duced into France. Although V. velutina inhabits tropical/subtropical
regions of Indo-China, it is predominantly a highland species. This
helps explain why V. velutina has spread rapidly in both South
Korea and France. The basic biology of V. velutina in South Korea

Table 1
The relative adults of hornet abundance in five locations in 2010 (see Fig. 1 for locations), compared to the native composition either where V. velutina is not present or during
1997–2001 in Busan city.

Busan city Haman-gun Masan city Gyeongsan city Yeongyang-gun

1997–2001 2010 2010 2010 2010 2010

V. velutina 0 454 (37%) 1 (5%) 4 (17%) 8 (16%) 0

V. simillima 924 (25%) 67 (5%) 0 0 2 (4%) 1 (4%)
V. analis 1101 (29%) 344 (28%) 2 (10%) 7 (29%) 12 (24%) 3 (11%)
V. crabro 609 (16%) 123 (10%) 16 (76%) 11 (46%) 20 (41%) 17 (63%)
V. mandarinia 1066 (29%) 233 (19%) 2 (10%) 2 (8%) 6 (12%) 5 (19%)
V. ducalis 33 (1%) 16 (1%) 0 0 1 (2%) 1 (4%)
V. dybowskii 3 (0.1%) 1 (0.1%) 0 0 0 0
Total 3736 1238 21 24 49 27

Please cite this article as: Choi, M.B., et al., Distribution, spread, and impact of the invasive hornet Vespa velutina in South Korea, J. Asia Pac.
Entomol. (2011), doi:10.1016/j.aspen.2011.11.004
4 M.B. Choi et al. / Journal of Asia-Pacific Entomology xxx (2011) xxx–xxx
Fig. 3. Effect of degree of urbanization and the invasion of V. velutina on the hornet composition in Busan city. (The numbers in bars are number of adults.)
appears to be similar to that found in its native range. In just 7 years,
V. velutina has become the most abundant species in the Busan region
by displacing hornets such as V. simillima that has a similar life cycle
and nesting preference. This is particularly noticeable in urban areas
where V. velutina represents over 70% of the hornet population. The
aggressive nature, large colony size, and preference for nesting in
urban areas mean that conflict with humans is inevitable. Currently,
119 Rescue Services emergency call-outs to deal with social wasp/
hornet problems are dominated by V. velutina (41%) and this number
is expected to increase. The rapid spread across France indicates that
V. velutina will continue to spread throughout South Korea and that it
will become an increasing problem as more beekeepers in rural areas
and people in urban areas become affected. Therefore, protection of
the public and beekeepers needs to go beyond the level of nest re-
moval to selecting preservation regions in which the variety of
wasps can be maintained within restricted areas to promote the coex-
istence of humans and wasps/hornets.
Acknowledgments
This research was supported by the Yeungnam University Post-
doctoral Program, 2011. We are thankful to Yeon-Tae Park and to
Table 2
The number of 119 emergency call outs in Geumjeong-gu, district in August to Septem-
ber 2010 to address social wasp or hornet problems.
Vespidae species
Vespa velutina
Polistes rothneyi
Vespa analis
Vespa simillima
Polistes nipponensis
Vespa crabro
Vespa mandarinia
Parapolybia indica
Vespula koreensis
total
Please cite this article as: Choi, M.B., et al., Distribution, spread, and impact of the invasive hornet Vespa velutina in South Korea, J. Asia Pac.
Entomol. (2011), doi:10.1016/j.aspen.2011.11.004
the chief and members of 119 Geumjeong Rescue Services (emergen-
cy call-outs) who provided samples of destroyed nests and adults in
Geumjeong-gu region of Busan city.
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Please cite this article as: Choi, M.B., et al., Distribution, spread, and impact of the invasive hornet Vespa velutina in South Korea, J. Asia Pac.
Entomol. (2011), doi:10.1016/j.aspen.2011.11.004