Journal of Stored Products Research 87 (2020) 101631

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Journal of Stored Products Research

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

Orientation of Tribolium castaneum (Herbst) (Coleoptera:

Tenebrionidae) adults at various distances to different concentrations
of aggregation pheromone 4,8-dimethyldecanal
D.M.S.K. Dissanayaka , A.M.P. Sammani , L.K.W. Wijayaratne *
Department of Plant Sciences, Faculty of Agriculture, Rajarata University of Sri Lanka, Puliyankulama, Anuradhapura, Sri Lanka

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

Article history: Tribolium castaneum is a serious pest of durable agricultural commodities during storage. The male
Received 30 March 2020 T. castaneum adults release the aggregation pheromone 4,8-dimethyldecanal (4,8 DMD). The 4,8 DMD is
Received in revised form produced commercially and available to be used in Dome traps along with their recommended kairo-
1 May 2020
mone but low trapping response is often reported. Concentration of 4,8 DMD influences the attraction of
Accepted 10 May 2020
T. castaneum adults but its intensity may vary under different warehouse settings. Further, the orien-
Available online 25 May 2020
tation response of T. castaneum adults from different distances to the synthetic 4,8 DMD is still uncertain.
Therefore, the objective of this research was to evaluate the effect of distance to the traps having
Keywords:
Tribolium castaneum
different 4,8 DMD concentrations on the orientation response of T. castaneum adults. The dome trap
Pheromone having two pheromone septa and kairomone was placed inside the experimental arena. From phero-
Kairomone mone, different concentrations were used. One-month-old T. castaneum adults were individually marked
Concentration and released from different distances at the same time. The adults reached the trap were counted 4 h
Distance following release. Alternatively the experiment was repeated without having the kairomone inside the
trap. One trap having neither the pheromone nor kairomone (empty trap alone) and one having hexane
only were used as controls. For all the pheromone concentrations used, the maximum trapping per-
centage was found when the beetles were released at 30 cm or 60 cm from the pheromone. Further, the
highest trapping percentage was given by 0.5 mL of 4,8 DMD. At a given distance, the traps having
pheromoneþkairomone better attract T. castaneum adults than those had only the pheromone. The study
concludes that the degree of attraction of T. castaneum adults varies with the distance from the trap and
the trap composition. Further studies are required to test the efficacy of 4,8 DMD and kairomones under
real warehouse settings.
© 2020 Elsevier Ltd. All rights reserved.

1. Introduction
Massive losses occurred during storage of agricultural produce is
a great challenge for the world food security as the global population
is estimated to be increased from current population of 7.6 billion to
9.8 billion by 2050 and 11.2 billion by 2100 (United Nations
Department of Economic and Social Affairs, 2017). Insect are great
respondents for these quantitative and qualitative losses occurred
throughout the world in different magnitudes (Wijayaratne et al.,
2018). Under storage conditions, the insect infestation and subse-
quent losses is reported in numerous types of food including cereal
* Corresponding author.
E-mail address: wollylk@yahoo.com (L.K.W. Wijayaratne).
grains, oil crops, pulses, processed food products, flour, confection-
aries and animal feed (Hagstrum and Subramanyam, 2006;
Hagstrum et al., 2012; Dissanayaka et al., 2018c; Sajeewani et al.,
2018, 2020; Wijayaratne et al., 2019; Wijerathne et al., 2020).
The management of insects infesting stored food is frequently
attended by the use of contact insecticides (Arthur et al., 2019),
fumigation (Hwaidi et al., 2017), high or low temperature exposure
(Beckett, 2011; Arthur et al., 2015). However, due to the limitations
faced by these practices such as phase out of fumigants such as
methyl bromide (Andersen et al., 2018), resistance development by
insects (Arthur et al., 1988; Opit et al., 2012), and negative impacts
on the biotic and abiotic environment (Fields, 1992; Arthur, 1996;
Phillips and Throne, 2010; Wijayaratne et al., 2018), biorational use
of pest management is encouraged (Phillips and Throne, 2010).
Tribolium castaneum (Herbst) (Coleoptera: Tenebrionidae) is a

https://doi.org/10.1016/j.jspr.2020.101631
0022-474X/© 2020 Elsevier Ltd. All rights reserved.
2 D.M.S.K. Dissanayaka et al. / Journal of Stored Products Research 87 (2020) 101631
major pest of food production lines and reported from almost all over
the world (Hagstrum and Subramanyam, 2006; Campbell et al.,
2010a,b. Precise monitoring and density determination are pre-
requisites for the management of this species (Semeao et al., 2013;
Arthur et al., 2014). The male T. castaneum adults biosynthesize and
release the aggregation pheromone 4,8, Dimethyldecanal (4,8 DMD)
from the setiferous glands or patches on the ventral side of the front
femora (Faustini et al., 1981, 1982). The 4,8 DMD attracts both sexes of
T. castaneum (Suzuki, 1980; Ming and Lewis, 2010). Different trap
types have been developed to monitor the population densities of
T. castaneum (Mullen,1992; Ho et al.,1997; Phillips et al., 2000; Fedina
and Lewis, 2007). The monitoring traps used for Tribolium species
have the aggregation pheromone 4,8 DMD and the commercially-
available kairomones (Campbell et al., 2002; Campbell et al.,
2010a,b; Dissanayaka et al., 2018a,b). Despite many studies con-
ducted on T. castaneum pheromone, information on beetle response
when used in commercial traps and under warehouse conditions is
limited (Campbell, 2012). The commercially-available traps often
contain kairomone along with the pheromone, and the detailed un-
derstanding on the beetle response to the traps is highly important for
efficient monitoring T. castaneum (Campbell, 2012; Dissanayaka et al.,
2020). Inadequate information on the trap density required for the
correct estimation of population (Buckman and Campbell, 2013)
limits the use of pheromone technology warranting further experi-
ments on the same.
Previous related studies on the behavioural response of
T. castaneum adults include use of electroantennography (EAG)
(Collins et al., 2007; Verheggen et al., 2007), pitfall bioassays
(Blotch Qazi et al., 1998; Seifelnasr et al., 1982), olfactometer bio-
assays (Romero et al., 2010). These studies, however, still lack
certain information required to achieve the maximum potential of
pheromone use. Such detailed studies on beetle availability around
the traps, effectiveness of both traps and attractants would enhance
the use of pheromone technology in food storage facilities
(Campbell, 2012). A previous study conducted in our laboratory has
reported the effective distance and pheromone concentration for
the attraction of T. castaneum adults to traps (Dissanayaka et al.,
2018a). However, that study had only the pheromone 4,8 DMD
inside traps whereas the commercial dome traps have kairomone
added with the pheromone. Such information on the attraction of
T. castaneum adults in the dome traps having both the pheromone
and kairomone is limited. Therefore the objectives of this experi-
ment were to determine the trapping efficiency of T. castaneum
adults located at different distances from the trap having phero-
mone and kairomone, and the effect of pheromone concentration
on the trapping of T. castaneum adults when only the pheromone or
both pheromone and kairomone are present in the traps.
2. Materials and methods
2.1. Insect cultures
The T. castaneum population used in the study was originally
collected from Nochchiyagama, Anuradhapura, Sri Lanka, and main-
tained in the laboratory since 2016. The T. castaneum cultures were
reared on wheat flour (whole meal flour) and maintained inside the
incubator (FH-1200, Hipoint Laboratory, Taiwan) at 30 ± 0.5  C,
65 ± 1% relative humidity and complete darkness. The parental adults
were introduced to 250 g flour medium, maintained for 15 days and
removed. Adults aged 30 days were used in the experiments.
2.2. Marking of adults
Adults were collected from the laboratory cultures and placed
on an Aluminum tray. The adults were brushed using paint brush
(No. 3) to remove any flour on adults. A small drop of nail polish
(Oem, Guangdong, China) was put on the elytra of the adults on the
mid dorsal line using a drawing pin. Different colors were used for
insects released at different distances (30 cm-white, 60 cm-blue,
90 cm-brown, 120 cm-red, 150 cm-grey and 180 cm-green). Later,
the painted adults were placed on a tissue paper to avoid spreading
of nail polish to other parts of the body and kept 3 h for drying.
2.3. Warehouse-release-recapture experiment
This experiment was conducted inside the warehouse
(12 m  10 m). The warehouse was cleaned using a biodegradable
disinfectant (Britol Disinfectant Pine, Antler Industries Pvt Ltd.,
Piliyandala, Sri Lanka), and any insects present were removed. The
experiment tested the trapping efficacy in a small arena neither
with the presence of food nor harbourage sites. Base of the exper-
imental arena was the cement floor, and edges covered with Teflon
(Polytetrafluoroethylene) (Sigma Aldrich, Saint Louis) to avoid in-
sects escaping from the arena (Fig. 1). Teflone was applied 10 min
before starting the experiments and repeated 1 h following the
commencement of experiment. All the experiments were con-
ducted between 9 am and 4 pm.
In this experiment, the effect of pheromone (4,8 DMD) dose (0.5,
1, 2, 3 or 4 mL) on the attraction of beetles was tested. The aggre-
gation pheromone (4,8 DMD) dissolved in hexane (1%) was applied
to the pheromone septa using a micropipette (Labnet International,
Inc., Poland). Along the midline of the long axis of experimental
arena, the Dome trap (Trece Inc., Adair, USA) having two phero-
mone septa (containing 4,8 DMD) was placed at 30 cm inside the
left edge (Fig. 1). The dome traps having the pheromone septa
containing hexane and just the empty trap (with neither phero-
mone nor kairomone) were used as controls. The Dome trap was
placed on the experimental arena 1 h before the experiment. On the
same midline where the trap was placed, six locations were marked
with 30 cm distance from each other. Twenty T. castaneum adults
marked with a particular colour, as described previously, were
placed on each location at the same time. After 4 h of releasing, the
adults trapped were collected, separated according to color code
and counted. The experiment was repeated for each pheromone
concentration used. From each pheromone concentration, four
replicate experiments were conducted. Following this, all the ex-
periments were repeated with the presence of pher-
omoneþkairomone (Trece Inc., Adair, USA) by adding 15 drops of
Kairomone (Trece Inc., Adair, USA). The actual temperature and
relative humidity during experiments were determined using
HOBO data loggers (Onset Computer Corporation, Bourne, MA). The
temperature and relative humidity in the experimental arena
during the experiment were 30 ± 1  C and 65 ± 0.5%, respectively.
2.4. Data analysis
The response variable was the T. castaneum adults reached the
trap following their release at different distances. The percentages
of T. castaneum adults reached the trap (having pheromone or
pheromoneþkairomones or the empty trap in the control)
following their release at a particular distance were transformed
using square root of the Arcsine value to accommodate the unequal
variances associated with the percentage data (Zar, 1999; Toews
et al., 2010; Burks and Kuenen, 2012; Trematerra et al., 2013;
Dissanayaka et al. 2018a,b; Wijayaratne and Rajapakse, 2018;
Dissanayaka et al., 2020). The normality of the data set was proven
by the Shapiro-Wilk test. The Bartlett’s test confirmed the homo-
geneity of variances. The transformed data were analyzed using
ANOVA of Statistical Analysis System (SAS Institute, 2002e2008)
and the means were separated by Tukey’s test. The significance
 D.M.S.K. Dissanayaka et al. / Journal of Stored Products Research 87 (2020) 101631
Fig. 1. Experimental arena on the floor of the warehouse showing the placement of trap and the edges covered with Teflon.
level was set at P ¼ 0.05.
3. Results
On the whole, the beetle attraction to trap varied with the dis-
tance from the trap (F5,156 ¼ 56.01; P < 0.0001) and pheromone
concentration (F6,156 ¼ 49.13; P < 0.0001). At 0.5 mL pheromone,
overall there was a significant difference in trap catch between
treatment (with pheromone) and control (without pheromone)
(F23,72 ¼ 95.31, P < 0.001) indicating that the pheromone (4,8 DMD)
attracted T. castaneum adults. Except at 30 and 60 cm, the trap catch
in the control at each of the other locations was zero for all the
pheromone concentrations tested (0.5, 1, 2, 3 and 4 mL) (Figs. 2 and
3). Further, at 0.5 mL pheromone concentration, the trap catch
significantly differed when the beetles were released at different
distances from the pheromone, in either situation when inside the
trap had the pheromone alone (F5,18 ¼ 29.70, P < 0.001) or pher-
omoneþkairomone (F5,18 ¼ 82.26, P < 0.001) (Fig. 2A).
When pheromone concentration was used at increased concen-
tration as 1 mL, overall, there was a significant difference in the trap-
ped adults percentages at different distances when only the
pheromone (F5,18 ¼ 13.72, P < 0.001) or both pheromoneþkairomone
was used (F5,18 ¼ 22.79, P < 0.001) (Fig. 2B). Use of 4,8 DMD at 2 mL also
demonstrated a pattern of beetle attraction similar to that with 1 mL.
There was a significant difference between trapped adults percentage
among the distances where the beetles were released. This occurred
with the pheromone alone (F5,18 ¼ 10.48, P < 0.001) and both pher-
omone and kairomone (F5,18 ¼ 18.06, P < 0.001) (Fig. 2C). When the
pheromone trap had 3 mL 4,8 DMD, trapping percentages among the
different distances significantly differed when only the pheromone
(F5,18 ¼ 13.56, P < 0.001) or both pheromone and kairomone
(F5,18 ¼ 15.19, P < 0.001) were present inside the trap (Fig. 3A). When
4 mL 4,8 DMD was used, there were significant differences among
trapping percentages for all the distances with the pheromone alone
(F5,18 ¼ 41.76, P < 0.001) or pheromoneþkairomone occasions
(F5,18 ¼ 21.44, P < 0.001) (Fig. 3B).
4. Discussion
The aggregation pheromone 4,8 DMD and kairomones are used
in monitoring devices to attract Tribolium species (Suzuki, 1980;
Campbell et al., 2002; Campbell et al., 2010a,b; Dissanayaka et al.,
2018a,b). Pheromone concentration and distance are a major fac-
tors that influences the response of T. castaneum adults (Elkinton and
Carder, 1984; Bell, 1984; Dissanayaka et al., 2018a). Dissanayaka
et al., 2018a reported that the maximum trapping of T. castaneum
3
is obtained until 60 cm from the trap when only the pheromone is
available inside the trap. This is again proven in the current study.
Campbell (2012) reported that maximum 40% of T. castaneum adults
attracted to Dome traps. Our study recorded higher trapping as 47%
with the pheromone alone whereas 56% of beetles released trapped
with pheromoneþkairomone condition. The current study also
proved that combination of pheromone and kairomone attracts
more adults than the pheromone alone as found in previous studies
(Phillips et al., 1993; Dissanayaka et al., 2018b). The response of
T. castaneum adults to pheromone mainly depends on qualitative
than the quantitative (Boake and Wade, 1984). In support of this
finding, in the current study, the trapping percentage was not
increased continuously when the pheromone concentration was
increased.
In contrast, our finding differs with some other previous find-
ings which report no difference in trapping by adding kairomone to
the system with pheromone (Willis and Roth, 1950; Romero et al.,
2010; Duehl et al., 2011). Hawkin et al. (2011) reported that the
trap catch of Tribolium confusum Jacquelin du Val, close relative of
T. castaneum, at 60 cm from the pheromone is low as 2%. In contrast,
the current study demonstrated much higher trapping of
T. castaneum adults at 60 cm from the pheromone when used in
different concentrations.
Dissanayaka et al. (2018b) reported that mee oil (Madhuca
longifolia) and coconut oil (Cocos nucifera) attract T. castaneum
adults similar to 4,8 DMD. They also reported that sunflower oil
(Helianthus annuus) and olive oil (Olea europaea) attract
T. castaneum adults similar to the commercial kairomones tested in
the current study. This also agrees with Phillips et al. (1993) that a
mixture of plant oils included in traps better attracts T. castaneum
than 4,8 DMD alone. Future research can be performed using these
oils in place of pheromone and/or kairomones used in this study.
Despite how the pheromone concentration and distance influ-
ence trapping of T. castaneum adults, the current study highlights
several limitations of use. These include the low trapping response
and reduced attraction at increased distances for which the future
research need to explore solutions. Further, the impact of phero-
mone concentration and distance on the trapping response of other
stored-product insects should be explored. Further, stored products
are damaged by lepidopteron species (Hill, 1990) and certain spe-
cies undergo diapause to pass the adverse environmental condi-
tions (Bell et al., 1983; Fields and Timlick, 2010; Wijayaratne and
Fields, 2012). As the physiological adaptations of these insects
differ from non-diapausing insects (Adkisson, 1966; Denlinger and
Lee, 2010), effects of pheromone concentration and distance on
trapping efficiency of those moth species would enhance the use of
4 D.M.S.K. Dissanayaka et al. / Journal of Stored Products Research 87 (2020) 101631

Fig. 2. Percentage (mean±SE) of Tribolium castaneum adults collected when released at different distances from the trap with or without the kairomone. The aggregation pher-
omone 4,8 DMD concentration varied as A: 0.5 mL, B: 1 mL, C: 2 mL. For a given 4,8 DMD concentration, the means followed by the same letter are not significantly different according
to Tukey’s test (P¼0.05).
 D.M.S.K. Dissanayaka et al. / Journal of Stored Products Research 87 (2020) 101631 5

Fig. 3. Percentage (mean±SE) of Tribolium castaneum adults collected when released at different distances from the trap with or without the kairomone. The aggregation pher-
omone 4,8 DMD concentration varied as A: 3 mL, B: 4 mL. For a given 4,8 DMD concentration, the means followed by the same letter are not significantly different according to
Tukey’s test (P¼0.05).

this technology for better protection of stored agricultural
commodities.
In general, biorational pest management claims a prominent
place in stored product protection (Phillips and Throne, 2010;
Wijayaratne et al., 2018). Pheromones are good candidates for bio-
rational pest management but accompanies limitations including
low trapping percentage as mentioned before. As combination of
treatments offer better control than use of a single treatment alone
(Banks, 1987; Banks and Fields, 1995), combined use of pheromone
with other promising biorational methods such as modified atmo-
sphere (Navarro et al., 2012; Wijayaratne et al., 2009); non-
ventilation/hermetic condition (Navarro et al., 2010; Hasaranga
et al., 2018; Wijayaratne et al., 2019); botanicals and kairomones
(Olivero-Verbel et al., 2013; Dissanayaka et al., 2018b; Stevens et al.,
2019), reduced-risk insecticides such as methoprene (Wijayaratne
and Fields, 2010; Arthur, 2016; Scheff et al., 2016, 2019) and spino-
sad (Hertlein et al., 2011; Kavallieratos et al., 2017; Wijayaratne and
Rajapakse, 2018) need to be explored. Future research need to test in
detail the reasons underlying the variation in the attraction of
T. castaneum adults in response to the traps having different pher-
omone concentrations and placed at different distances. The
developed technology also need to be investigated under complex
settings at grain storage and food processing facilities to determine
the required density and composition of traps. Testing these avenues
following a proper design would ensure successful pest manage-
ment in stored agricultural products while ensuring the safety of
biotic and abiotic environment.
6 D.M.S.K. Dissanayaka et al. / Journal of Stored Products Research 87 (2020) 101631
Declaration of competing interest
The authors declare that they have no known competing
financial interests or personal relationships that could have
appeared to influence the work reported in this paper.
CRediT authorship contribution statement
D.M.S.K. Dissanayaka: Conceptualization, Investigation, Data
curation, Formal analysis, Writing - original draft. A.M.P. Sammani:
Investigation. L.K.W. Wijayaratne: Conceptualization, Methodol-
ogy, Validation, Supervision, Writing - review & editing, Resources,
Funding acquisition, Project administration.
Acknowledgements
The authors are grateful for the Sri Lanka Council for Agricul-
tural Research Policy (SLCARP) (Grant No: NARP/16/RUSL/AG/01)
for financial assistance provided.
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