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Impacts of 2 species of predatory Reduviidae on bagworms in oil palm


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DOI: 10.1111/1744-7917.12309

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Insect Science (2016) 00, 1–10, DOI 10.1111/1744-7917.12309

ORIGINAL ARTICLE

Impacts of 2 species of predatory Reduviidae on bagworms


in oil palm plantations

Syari Jamian1 , Ahmad Norhisham2 , Amal Ghazali3 , Azlina Zakaria4 and Badrul Azhar3,5
1 Faculty of Agriculture, Universiti Putra Malaysia, Serdang, Selangor, Malaysia; 2 Cardiff School of Biosciences, Cardiff University, Cardiff,
Wales, UK; 3 Faculty of Forestry, Universiti Putra Malaysia, Serdang, Selangor, Malaysia; 4 Sime Darby Research Sdn. Bhd., Banting,
Selangor, Malaysia and 5 Biodiversity Unit, Institute of Bioscience, Universiti Putra Malaysia, Serdang, Selangor, Malaysia

Abstract Integrated pest management (IPM) is widely practiced in commercial oil palm
agriculture. This management system is intended to minimize the number of attacks by pest
insects such as bagworms on crops, as well as curb economic loss with less dependency
on chemical pesticides. One practice in IPM is the use of biological control agents such
as predatory insects. In this study, we assessed the response of predatory natural enemies
to pest outbreak and water stress, and document the habitat associations of potential pest
predators. The abundances of 2 predatory insect species, namely Sycanus dichotomus and
Cosmolestes picticeps (Hemiptera: Reduviidae), were compared bagworm outbreak sites
and nonoutbreak sites within oil palm plantations. We also examined habitat characteristics
that influence the abundances of both predatory species. We found that the abundance of
C. picticeps was significantly higher in bagworm outbreak sites than in nonoutbreak sites.
There were no significant differences in the abundance of S. dichotomus among outbreak
and non-outbreak sites. Both species responded negatively to water stress in oil palm
plantations. Concerning the relationship between predatory insect abundance and in situ
habitat quality characteristics, our models explained 46.36% of variation for C. picticeps
and 23.17% of variation for S. dichotomus. Both species of predatory insects thrived from
the planting of multiple beneficial plants in oil palm plantations. The results suggest that
C. picticeps can be used as a biological agent to control bagworm populations in oil palm
plantations, but S. dichotomus has no or little potential for such ecosystem service.
Key words bagworms; beneficial plants; biological control; habitat quality; integrated
pest management; oil palm; predatory insects

Introduction 55% between 2001 and 2006 (see http://faostat.fao.org).


Due to this, commercial oil palm cultivation has
Rising global demand for palm oil has promoted oil palm transformed many areas of arable lands into oil palm
(Elaeis guineensis) as the world’s most rapidly expanding plantation. In Malaysia alone, the area planted with oil
crop (Laurance et al., 2010). Oil palm provides an palm has increased from 1.8 million ha to 5.2 million ha
efficient and inexpensive resource for vegetable oil and between 1990 and 2013 (see http://www.mpob.gov.my).
biofuel in the world market today (Fitzherbert et al., 2008; As the second largest producer of palm oil after Indonesia,
Lam et al., 2009), with an increase in global production of Malaysia contributed to 80.5% of global production, with
an export value of US$6.3 billion in 2004 (Koh & Wilcove,
2007). Apart from its importance for economy and food
Correspondence: Badrul Azhar, Biodiversity Unit, Institute security, oil palm plantations also show farmland biodi-
of Bioscience, Universiti Putra Malaysia, Serdang, Selangor versity in terms of flora and fauna (Rice & Greenberg,
43400, Malaysia. Tel: +603 8946 7206; fax: +603 8943 2514; 2000), although this may be limited to a small number of
email: b_azhar@upm.edu.my species (Turner & Foster, 2009; Azhar et al., 2011).
1

C 2015 Institute of Zoology, Chinese Academy of Sciences
2 S. Jamian et al.

With the increasing rate of oil palm expansion, insect by introducing flowering plants at the edges of oil palm
pest infestation such as bagworms (such as Metisa plana plantations (Wilcove & Koh, 2010). However, there is no
Walker and Pteroma pendula Joannis), nettle caterpillars clear evidence that the presence of beneficial plants im-
(Setora nitens), and the rhinoceros beetle (Oryctes spp.) proves the population abundance of predatory insects in
in oil palm plantation has become a primary concern oil palm plantations. Previous studies have shown that the
among stakeholders (Foster et al., 2011). Bagworms have presence of beneficial flowering plants in tropical agroe-
been recognized as the most destructive insect pest in the cosystems can increase the abundance of natural enemies,
oil palm plantations of Peninsular Malaysia (Basri et al., including parasitoids (Wratten et al., 2002). A study by
2001; Rhainds et al., 2002; Norman & Basri, 2010). Basri et al. (1999) reported that flowering plants may pro-
Infestations of early instar larvae cause defoliation, long adult parasitoid life spans. Efforts are being made to
becoming worse as the larvae grow. Larger populations quantify the usefulness of nectariferous plants in encour-
cause more extensive damage and infestations become aging natural predatory insect populations, but results are
more intense in oil palm monocultures (Ho, 2002). thus far inconsistent (Ho & Teh, 1999). Little is known
The use of chemical pesticides has proven to be about the response of predatory insect populations to fluc-
ineffective as they promote pest resistance and eliminates tuations in potential prey species. It is thus important to
beneficial insects such as pollinators and predators determine the benefits of flowering plants for predatory
(Cheong et al., 2010). Predatory insects are known to be insects so that efforts maybe more focused on their use in
an excellent tool for the biological control of insect pests oil palm plantations.
(Pimentel et al., 1992; Cardinale et al., 2003; Fazal et al., Maintenance of predatory insect populations as biolog-
2012). Insects such as parasitoids and hemipterans serve ical control agents requires assessment of habitat quality
an important role in regulating bagworm and caterpillar to distinguish which characteristics support higher pop-
populations in oil palm plantations (Basri et al., 1995; ulations of predatory insects. Thus, this study proposes
Norman & Basri, 2007; Cheong et al., 2010). Currently, 6 the 3 following research questions to investigate the pop-
hemipteran species have been reported as potential preda- ulation response and habitat associations of predatory in-
tors in oil palm plantations in Peninsular Malaysia, namely sects in oil palm plantations: (i) how do the abundances of
Cosmolestes picticeps (Yusdayati, 2008), Callimerus predatory insects respond to non-outbreak and bagworm
arcufer (Basri et al., 1996), Sycanus dichotomus (Jamian outbreak areas? We predicted that predatory insect abun-
et al., 2011), Andrallus spinidens, Platynopus melachan- dance might be greater in bagworm outbreak areas than
tus, and Cantheconidea furcellata (Khoo & Chan, 2000). in nonoutbreak areas because the population outbreaks
Integrated pest management (IPM) is a series of of bagworms would increase the abundance of predatory
essential strategies which provide a long-term prevention insects; (ii) how do the abundances of predatory insects
of pests while limiting environmental damage (Kogan, respond to water stress? We predicted that predatory insect
1998). IPM systems keep pest populations below abundance might be lower at sites without standing water;
damaging levels via insects or other animal and plant and (iii) to what extent does habitat quality characteristics
management tactics with minimal environmental impacts influence predator abundance in oil palm plantations? We
(Romoser & Stoffolano, 1998; Wood, 2002). Successful predicted that predatory insect abundance might be influ-
implementation of IPM systems in oil palm plantations enced by aspects of particular vegetation structures, such
has been reported by Caudwell (2000). Preserving as ground vegetation coverage and number of beneficial
beneficial and nontarget organisms is an important part plant species.
of the system. However, propagation and establishment
of natural enemies such as predators and parasitoids
are dependent on flowering plants, as flowering plants Materials and methods
support higher biodiversity and provide an important
source for insect foraging activity (Donald, 2004; Ellis Study site
et al., 2005). With regard to this, to increase predatory
insect populations such as C. picticeps and S. dichotomus, We surveyed 4 oil palm plantations, namely South-
beneficial plants such as Turnera subulata, Cassia coba- ern Perak estate (3797.27 ha) in the state of Perak
nensis, Antigonon leptopus, and Euphorbia heterophylla (N 03°48 45.09 , E 101°00 56.01 ), Bukit Talang estate
are commonly planted alongside roads within oil palm (4 428 ha) in the state of Selangor (N 03°23 58.4 , E
plantations. 101°18 94.2 ), Bukit Senorang estate (1628.10 ha), and
Diversity of crop plants and proper strategies can influ- South East Pahang estate in the state of Pahang (N 03°07
ence species richness in agricultural landscapes (Donald, 04.3 , E 102°24 48.0 and N 03°05 23.5 , E 102°24
2004). Populations of predatory insects maybe increased 53.7 ) (Fig. 1). Plantations were defined as large-scale oil

C 2015 Institute of Zoology, Chinese Academy of Sciences, 00, 1–10
Predatory insects in oil palm agriculture 3

and visual identification. Sweep netting was standardized


using a set number of back-and-forth sweeps (between 10
and 20) approximately 1 m in length taken every other
pace while walking at a consistent speed through the veg-
etation (Ausden & Drake, 2006). Predator species were
identified based on their morphological characteristics,
while the number of each species was recorded. This study
was carried out from September 2013 to Jun 2014.

Water stress, vegetation characteristics, and beneficial


plants

Water stress was determined based on the presence


or absence of standing water on sites. Standing water
was characterized as any man-made source of water of
Fig. 1 Map showing location of the study areas and 100 tran- greater than 50 cm depth, including ponds, drain wa-
sects in the states of Perak (A), Selangor (B), and Pahang (C). ter, and reservoirs. We measured and counted 4 local
habitat quality characteristics at each transect. We com-
pared these factors among outbreak and nonoutbreak sites
palm cultivation areas covering more than 50 ha each, and
as well as among sites with standing water and with-
were managed by private businesses (Azhar et al., 2011).
out standing water. We counted the number of beneficial
To maintain high yield of oil palm fruit bunch, agrochem-
plants planted by plantation management. The coverage
icals such as pesticides and fertilizers are commonly used
of ground vegetation was estimated in a rectangle plot of
by managers in oil palm plantations. In addition, conven-
100 m2 (length × width = 100 m × 1 m) on all tran-
tional plantations are characterized by modern facilities
sects. The height of ground vegetation was measured,
and infrastructure, such as paved roads, perimeter fences,
taking 4 measurements per transect. In terms of floristic
worker settlements and mills.
composition, ground vegetation cover comprised benefi-
Outbreak sites were defined by the presence of at least
cial plants (e.g., Antigonon leptopus, Turnera subulata,
10 bagworm larvae per frond (Wood, 2002). This is the
Asystasia gigantica, Cassia cobanensis, and Euphorbia
pest population density at which control measures should
heterophylla), grasses (e.g., Paspalum conjugatum and
be initiated to prevent an increasing pest population from
Centosteca lappacea), broadleaves (e.g., Croton hirtus
reaching the level of economic injury (Stern et al., 1973).
and Mikania micrantha), ferns (e.g., Nephrolepis biser-
Southern Perak and South East Pahang estates were identi-
rata and Dicnopteris linearis), and woody shrubs (e.g.,
fied as outbreak sites. These estates were typically sprayed
Clidemia hirta, Lantana camara, and Melastoma mala-
with chemical pesticides (i.e., Cypermethrin) to control
bathricum). The number of beneficial plant species at
bagworm populations. However, there was no insecticide
each transect was recorded.
intervention throughout this study period.
Oil palms may grow to heights of 18 m or greater. The
trunks of young and mature palms are overgrown with
Insect sampling epiphytes. On the other hand, the older palms are charac-
terized by smoother trunks. We used a laser rangefinder
We conducted sampling on clear days (without rains to determine the height of palm trees. Height measure-
or heavy clouds) from 0700 to 1100 h and again from ment was determined from the base of oil palm to the top
1500 to 1800 h by traversing slowly along straight line- (base of fronds). The height of oil palm stand was taken
transects that spanned 100 m. We established 100 tran- at intervals of 25 m. Water stress was determined by the
sects alongside plantation roads, with 50 transects at presence or absence of standing water at study sites. This
each treatment (outbreak sites versus nonoutbreak sites). categorical variable was determined in the surrounding
Each transect was spaced 100 m apart to ensure inde- area of transect (within the radius of 10 m).
pendence of observations. We surveyed each transect
3 times. There were 40, 10, 10, and 40 transects at South- Statistical analysis
ern Perak estate, Bukit Senorang estate, South East Pa-
hang estate and Bukit Talang estate, respectively (Fig. 1). We performed a Shapiro–Wilk’s test to check for nor-
To sample adult predatory insects, we used a sweep net mality in the abundance of predatory insects. Abundance


C 2015 Institute of Zoology, Chinese Academy of Sciences, 00, 1–10
4 S. Jamian et al.

Table 1 Statistical summary of habitat variables that were measured or counted from 100 transects (50 in each type of site) and each
repeated 3 times. One-way ANOVA was used to compare the variables between outbreak sites and nonoutbreak sites. Visit was used as
a block.

Outbreak site Nonoutbreak site


Habitat variable F value P value
Mean SD Mean SD

Ground vegetation coverage (%) 77.33 15.40 52.25 22.61 96.67 <0.001
Height of ground vegetation (cm) 128.6 21.39 85.04 27.60 227.12 <0.001
Height of oil palm stand (m) 3.388 1.004 6.414 1.430 313.82 <0.001
Richness of beneficial plant 2.373 0.909 1.78 0.850 36.09 <0.001

data were square-root transformed prior to inferential oil palm stand height, differed significantly among sites
analysis. To answer our first and second hypothesis, we with and without standing water (Table 2).
performed one-way ANOVA to compare the abundance of
predatory insects among bagworm outbreak and nonout-
break plantations (Zar, 1999; Quinn & Keough, 2002). Predatory insect populations in outbreak sites
We used different visits as blocks in the analysis. We and nonoutbreak sites
also used unbalanced design ANOVA to compare the ef-
fects of different times of day (morning survey = 191 We recorded a total of 1230 individuals belong-
times; afternoon survey = 109 times) and water availabil- ing to 2 species of predatory insects, namely Cosmo-
ity (absence of water = 126 times; presence of water = lestes picticeps (n = 1045) and Sycanus dichotomus
174 times) on predator abundance. The means of abun- (n = 185). As expected, we found that the abundance
dance were predicted using regression models. To test for of C. picticeps was significantly higher (df = 1; F =
the interaction between water availability and outbreak 28.49; P < 0.001) in bagworm outbreak sites (mean ±
status, we performed a two-way ANOVA test. SE = 4.96 ± 0.472 individuals) than nonoutbreak sites
To address the second hypothesis, we used General (mean ± SE = 2.007 ± 0.240 individuals) (Fig. 2). Dif-
Linear Models (GLMs) (Schall, 1991). We developed ferent visits had no effect on the species abundance (df =
a predictive model for each species of predatory insect. 2; F = 1.35; P = 0.261). However, our results revealed
In each model, insect abundance was used as dependent that the abundance of S. dichotomus (df = 1; F = 2.34;
variable. The independent variables were richness of ben- P = 0.127) in bagworm outbreak sites (mean ± SE =
eficial plants, mean height of ground vegetation, mean 0.673 ± 0.0833 individuals) did not differ significantly
height of oil palm, and coverage of ground vegetation. with nonoutbreak sites (mean ± SE = 0.56 ± 0.0919 indi-
To deal with collinearity, we removed predictor variables viduals) (Fig. 2). The species abundance was significantly
from the models if correlations were strong (|r|> 0.7, affected by different visits (df = 2; F = 20.95; P < 0.001).
where r is the coefficient of correlation) (Dormann et al., The abundances of C. picticeps and S. dichotomus were
2013). We used Poisson distribution and log-link function not influenced by different times of day (df = 1; F =
to fit the data. Visit month was used as an absorbing 2.83; P = 0.094 and df = 1; F = 2.90; P = 0.090, re-
factor in the analysis. The best models were selected spectively). Our analysis indicated that the presence of
based on R2 . All statistical analyses were conducted in standing water was important to C. picticeps (df = 1; F =
GenStat 12 (VSNI Hemel, Hempstead, UK). 178.01; P < 0.001). The abundance of C. picticeps was
greater at sites with no water stress (mean ± SE = 4.052 ±
0.006 individuals) than at those with water stress (mean ±
Results SE = 0.160 ± 0.008 individuals). Similarly, the abun-
dance of S. dichotomus was significantly greater (df =
Habitat quality at sites with different levels of outbreak 1; F = 176.97; P < 0.001) at sites with no water stress
status and water availability (mean ± SE = 0.612 ± 0.001 individuals) than those with
water stress (mean ± SE = 0.004 ± 0.002 individuals).
There was a significant difference in each habitat The interaction between the explanatory variables
variable among outbreak sites and non-outbreak sites water availability and outbreak status was significant
(Table 1). All habitat variables, with the exception of (df = 1; F = 18.51; P < 0.001); hence, that water


C 2015 Institute of Zoology, Chinese Academy of Sciences, 00, 1–10
Predatory insects in oil palm agriculture 5

Table 2 Statistical summary of habitat variables that were measured or counted from 126 dried sites and 174 water-available sites.
Unbalanced design ANOVA was used to compare the variables between sites with standing water and without standing water. Visit was
used as a block.

Dried site Water-available site


Habitat variable F value P value
Mean SD Mean SD

Ground vegetation coverage (%) 64.80 11.82 64.79 28.57 5.72 0.017†
Height of ground vegetation (cm) 88.97 31.94 119.7 27.11 91.13 <0.001‡
Height of oil palm stand (m) 5.085 2.264 4.768 1.689 0.06 0.814§
Richness of beneficial plant 1.373 0.517 2.586 0.820 224.36 <0.001¶

Predicted means ± SE for site without standing water and site with standing water are 63.68% ± 1.047% and 55.21% ± 1.039%,
respectively (computed from regression model).

Predicted means ± SE for site without standing water and site with standing water are 82.98% ± 1.027% and 116.68% ± 1.023%,
respectively (computed from regression model).
§
Predicted means ± SE for site without standing water and site with standing water are 4.496 ± 1.043 m and 4.438 ± 1.036 m,
respectively (computed from regression model).

Predicted means ± SE for site without standing water and site with standing water are 1.332 ± 0.0005 and 2.509 ± 0.0003, respectively
(computed from regression model).

availability effects on the abundances of C. picticeps


were not the same for both outbreak status levels, and
outbreak status effects on the abundances of C. picticeps
were not the same for the 2 levels of water availability. In
contrast, the interaction between the explanatory variables
water availability and outbreak status was not significant
for S. dichotomus (df = 1; F = 3.65; P = 0.057).

Important habitat quality characteristics for determining


predatory insect abundance

For C. picticeps, our model explained 46.36% of vari-


ations in the data. All predictor variables were important
for maintaining populations of C. picticeps in oil palm
plantations (Fig. 3). Specifically, we found that the abun-
dance of C. picticeps increased with increasing height
of ground vegetation (slope = 0.02125; Wald = 261.1;
P < 0.001). Richness of beneficial plants increased the
abundance of C. picticeps (slope = 0.3668; Wald = 103.4;
P < 0.001). Site with tall oil palm had lower C. picticeps
abundance (slope = -0.1354; Wald = 57.1; P < 0.001).
The abundance of C. picticeps increased with increasing
ground vegetation coverage (slope = 0.00860; Wald =
24.7; P < 0.001).
Fig. 2 Abundances of Sycanus dichotomus were similar in non- The second model accounted for 23.17% of the
outbreak and outbreak sites. In contrast, abundances of Cosmo- variations that explained the relationships between S. di-
lestes picticeps differed significantly among non-outbreak and chotomus and predictor variables. Similar to C. picticeps,
outbreak sites. population levels of S. dichotomus in oil palm plantations
were determined by 4 predictor variables (Fig. 3). We


C 2015 Institute of Zoology, Chinese Academy of Sciences, 00, 1–10
6 S. Jamian et al.

Fig. 3 Responses of Sycanus dichotomus and Cosmolestes picticeps to multiple vegetation structure characteristics. Both predatory
insects require similar habitat qualities in oil palm plantations.


C 2015 Institute of Zoology, Chinese Academy of Sciences, 00, 1–10
Predatory insects in oil palm agriculture 7

found that the richness of beneficial plants increased the most critical aspects of oil palm plantation management.
abundance of S. dichotomus (slope = 0.4538; Wald = Oil palms require a lot of water to ensure good growth
43.71; P < 0.001). Similar to C. picticeps, the abundance and fruit yield; hence, an unstable supply of water can
of S. dichotomus decreased with increasing height of oil create unnecessary stress to the oil palms and adversely
palm (slope = -0.3399; Wald = 31.97; P < 0.001). The affect their productivity. Moreover, pest insects may attack
abundance of S. dichotomus increased with increasing drought-stressed oil palms.
height of ground vegetation (slope = 0.004056; Wald
= 10.73; P = 0.001). We also found that the abundance
of S. dichotomus increased with increasing ground Effects of in situ habitat quality characteristics on
vegetation coverage (slope = 0.002089; Wald = 6.42; predatory insect abundance
P = 0.011).
Our results have indicated that the abundances of C. pic-
ticeps and S. dichotomus increased with the richness of
Discussion beneficial plants. The findings seem to be consistent with
other research which found beneficial flowering plants
Responses of predatory insect populations to bagworm such as C. cobanensis and T. subulata are able to attract
infestations natural enemies of bagworm (Ho, 2002; Yusdayati et al.,
2014). These host plants provide refuge and nectar for nat-
Our study has indicated that abundance of C. picti- ural enemies. By planting various beneficial plants in oil
ceps was higher in bagworm outbreak sites than non- palm plantations, oil palm growers will be less dependent
outbreak sites. There was also no significant difference in on chemical pesticides to control bagworm populations.
S. dichotomus among bagworm outbreak sites and non- Because some of the beneficial plants are not perennial
outbreak sites. This reflects the fact that C. picticeps is species (e.g., C. cobanensis), such plants require continu-
a major predatory insect for bagworm infestation in oil ous replanting. Alternative host plants (e.g., T. subulata)
palm plantations. Similar results were also discussed in a should be planted as well during the absence of annual
study by Cheong et al. (2010), the C. picticeps proved to species (Yusdayati et al., 2014).
be a significant predator of bagworms as it was observed The abundances of C. picticeps and S. dichotomus de-
attacking P. pendula. creased with height of oil palm stands. It is likely that
Both species of predatory insects responded negatively some individuals were captured around ground vegetation
to water stress. This stress occurred due to the absence while others escaped into oil palm canopy. Similar results
of standing water in oil palm plantations. The natural have been shown in a study conducted by Pierre and Idris
source of water in most plantations is typically rainfall. (2013), in which the weaver ant, Oecophylla smaragdina,
The demand for water may exceed the available amount was used instead of hemipterans to reduce bagworm infes-
during a certain period or when poor quality restricts its tation. Based on their results, field surveys carried out on
use. Water stress may affect the growth of the plants, in- oil palm plantations in Peninsular Malaysia have shown
directly affecting the insect community. Hence, rainfall the presence of ants in short and young palms. This is
might influence the abundances of both C. picticeps and also supported by a study (Lowman, 1985), in which trees
S. dichotomus. Our analysis has confirmed that the avail- with young leaf situated closer to the ground experienced
ability of standing water in oil palm plantations is vital for a higher herbivory activity by insects in the rainforest of
predatory insects. Similar results were shown in a study Australia. In addition, both predatory insects mostly con-
by Zhu et al. (2014), in which changes in the amount trol bagworm populations in shorter oil palms. Height of
of rainfall significantly affected the diversity, abundance, oil palm tree indeed plays a major role in determining the
and trophic structure of the insect community in the stud- abundance of predatory insects in oil palm plantations.
ied area. The abundance of C. picticeps was significantly This study also indicates that shorter oil palms have a
lower at water stressed sites as predicted. The results are greater chance of surviving bagworm infestation, as the
also consistent with studies conducted by Cheong et al. abundance of predatory hemipterans was higher near such
(2010) and Chung and Sim (1991), in which the num- trees than the taller oil palms.
ber of outbreak cases of bagworms showed a significant Both hemipteran species also show increased abun-
increase in a relatively high rainfall period and not nec- dance with increasing coverage of ground vegetation.
essarily in dry months. Greater amount of food equals a Ground vegetation cover can provide predatory insects
greater number of predatory insects, as their prey resource with alternative sources of food, shelter, and oviposition
is plentiful. Nonetheless, water management is one of the sites (McEwan et al., 2005). Therefore, ground vegetation


C 2015 Institute of Zoology, Chinese Academy of Sciences, 00, 1–10
8 S. Jamian et al.

cover is essential for maintaining predatory insect abun- Azhar, B., Lindenmayer, D.B., Wood, J., Fischer, J., Manning,
dance. Habitat quality characteristics such as ground veg- A., McElhinny, C. and Zakaria, M. (2011) The conservation
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of beneficial plant species influence the predatory insect peat swamp forest for birds. Forest Ecology and Management,
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for maintaining a steady number of predatory insects in emies of bagworms, Metisa plana (Lepidoptera: Psychidae)
the plantations. and their impact on host population regulation. Crop Protec-
Our study is the first to have investigated the responses tion, 14, 637–645.
of predatory insects to bagworm outbreak and water Basri, M.W., Ramlah Ali, A.S., Moslim, R. and Arshad, O.
stress in oil palm plantations. We also examined habitat (1996) Biological efficacy of three commercial products of
characteristics, particularly vegetation structures, which Bacillus thuringiensis for the control of bagworms, Metisa
may be important for maintaining the populations of plana and Mahasena corbetti (Lepidoptera: Psychidae) on oil
predatory insects in these plantations. The study find- palm. PORIM International Palm Oil Congress. Competitive-
ings have improved understanding of predatory insects ness for the 21st Century (ed. D. Ariffin), pp. 369–378. Palm
and their role for keeping oil palm plantations free from Oil Research Institute of Malaysia, Malaysia.
bagworm outbreaks. Our findings provide new insights Basri, M.W., Siburat, S., Ravigedari, S. and Othman, A. (1999)
into how to maintain predatory insect levels in oil palm Beneficial plants for the natural enemies of the bagworm
plantations. In order to use natural predatory insects as in oil palm plantations. Proceeding of the 1999 PORIM In-
biological control of pest insect populations, the main- ternational Palm Oil Congress: Emerging Technologies and
tenance of local habitat complexity and farmland biodi- Opportunities in the Next Millennium. Palm Oil Research In-
versity are is in plantation management as well as IPM. stitute of Malaysia, Malaysia, 165–179.
Stakeholders should manage oil palm plantations so that Basri, M.W., Norman, K. and Othman, A. (2001) Field impact
the plantations become more favorable to agents of bio- of beneficial plants on the parasitism levels of the bagworm,
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cultural Conference 2001 International Palm Oil Congress
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Acknowledgments
Malaysian Palm Oil Board.
Cardinale, B.J., Harvey, C.T., Gross, K. and Ives, A.R. (2003)
We wish to thank A. Jambari, N. L. Ibrahim, and M.
Biodiversity and biocontrol: emergent impacts of a multi-
Syafiq from Universiti Putra Malaysia for assisting this
enemy assemblage on pest suppression and crop yield in an
project at the study sites. This project was supported by
agro-ecosystem. Ecology Letters, 6, 857–865.
the RUGS Initiative 6 scheme (Project no. 9327100). We
Cheong, Y.L., Sajap, A.S., Hafidzi, M.N., Omar, D. and Abood,
are also grateful to the estate managers and staffs of Sime
F. (2010) Outbreaks of bagworms and their natural enemies in
Darby Plantation, Southern Perak, and United Malacca
an oil palm, Elaeis guineensis, plantation at Hutan Melintang,
Berhad for permitting us to access study sites within the
Perak, Malaysia. Journal of Entomology, 7, 141–151.
oil palm estates. S. Jamian and B. Azhar conceived the
Chung, G.F. and Sim, S.C. (1991) Bagworm Census and Control:
research idea, designed the study, and performed analyses.
A Case Study. PORIM, Kuala Lumpur, 4, 433–442.
S. Jamian collected data at the study sites. All authors
Caudwell, R.W. (2000) The successful development and im-
discussed the study results and wrote the manuscript. We
plementation of an integrated pest management system for
also thank J. Vafidis for revising the final draft.
oil palm in Papua New Guinea. Integrated Pest Management
Reviews, 5, 297–301.
Disclosure Davis, A.J., Holloway, J.D., Huijbregts, H., Krikken, J., Kirk-
Spriggs, A.H. and Sutton, S.L. (2001) Dung beetles as indi-
The authors have declared that no competing interests cators of change in the forests of northern Borneo. Journal of
exist. Applied Ecology, 38, 593–616.
Donald, P.F. (2004) Biodiversity impacts of some agricultural
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C 2015 Institute of Zoology, Chinese Academy of Sciences, 00, 1–10

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