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Interference of Plant Essential Oils on the Foraging Behavior of
Solenopsis invicta (Hymenoptera: Formicidae)
Author(s): Jie Wang, Xiaolong Qiu, Ling Zeng and Yijuan Xu
Source: Florida Entomologist, 97(2):454-460.
Published By: Florida Entomological Society
DOI: http://dx.doi.org/10.1653/024.097.0215
URL: http://www.bioone.org/doi/full/10.1653/024.097.0215
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454 Florida Entomologist 97(2) June 2014
INTERFERENCE OF PLANT ESSENTIAL OILS ON THE FORAGING
BEHAVIOR OF SOLENOPSIS INVICTA (HYMENOPTERA: FORMICIDAE)
Jie Wang, Xiaolong Qiu, ling Zeng and YiJuan Xu*
Red Imported Fire Ant Research Center, South China Agricultural University, Guangzhou 510642, China
*Corresponding author’s; E-mail: xuyijuan@yahoo.com
abstract
Plant essential oils restrained the course of foraging behavior of individual worker ants
and also influenced worker recruitment and food transport. Worker search times increased
significantly among the plant oil treatments. The longest search time was observed for ants
treated with essential oils of Capsicum annuum L. (Solanales: Solanaceae) and Cedrus deo-
dara (Roxb.) G.Don (Pinales: Pinaceae). Different essential oils have significantly different
effects on the recruitment of the workers, the amount of food transported, and the time
spent foraging.
Key Words: fire ants, plant essential oil, foraging behavior, interfering, repellency ef-
fects
resumen
Los aceites esenciales de plantas restringieron la dirección de la conducta individual de
forrajeo de individuos de las hormigas obreras y también influyeron la selección de los tra-
bajadores y el transporte de alimentos. El tiempo de búsqueda de los trabajadores incre-
mentaron significativamente entre los tratamientos con aceites vegetales. Se observó que
el mayor tiempo de búsqueda en las hormigas tratadas con aceites esenciales de Capsicum
annuum L. (Solanales: Solanaceae) y Cedrus deodara (Roxb.) G.Don (Pinales: Pinaceae).
Diferentes aceites esenciales tienen efectos significativamente diferentes en la seleción de
los trabajadores, la cantidad de alimentos transportados, y el tiempo de forrajeo.
Palabras Clave: hormigas de fuego, aceite esencial de la planta, comportamiento de forrajeo,
interferir, efectos de repelencia
The red imported fire ant, Solenopsis invicta
Buren (Hymenoptera: Formicidae) causes seri-
ous damage to humans, animals and the environ-
ment. The red imported fire ant was discovered in
Southern China at the end of 2004. The ant has in-
vaded schools, rice fields, lawns, and other public
areas in both cities and rural locations (Zeng et al.
2005). Chemical treatments have been adopted as
the main method for the control of this important
medical and agricultural pest (Lofgren et al. 1975;
Adams et al. 1983; Lofgren 1986; Adams et al. 1988;
Drees & Gold 2003). However, in view of the envi-
ronmental pollution resulting from such chemicals,
the use of many types of pesticides is prohibited in
areas within the vicinity of food or water sources
(Williams & DeShazo 2004). Therefore, there is a
growing interest in treatment methods that are
non-toxic or only slightly toxic in the environment.
This interest includes fire ant repellents that can
potentially be applied in schools, nursing homes,
hospitals and other pesticide-sensitive sites. The
application of fire ant repellents to quarantined
nurseries and equipment used for working the soil
could also play an additional role in preventing
the spread of fire ants to non-infested areas (Chen
2009).
Previous reports confirmed that a number of ma-
terials and compounds were effective repellents of
fire ants (Oi & Williams 1996; Vander Meer et al.
1998; Anderson et al. 2002; Vogt et al. 2002; Chen
2005). Certain landscaping materials, such as cedar
and cypress mulch, are known to be effective repel-
lents of fire ants (Thorvilson & Rudd 2001). In addi-
tion, a recent study in Texas (USA) tested whether
a certain type of Old World bluestem grass, ‘WW-B
Dahl’ could effectively repel the red imported fire
ant (Sternberg et al. 2006). The results of that study
showed that the number of fire ant mounds were
much lower in WW-B Dahl pastures than in fields
planted with other types of grass. Furthermore,
WW-B Dahl grass proved to be an attractive alter-
native to ranchers and farmers because this grass is
favored by cattle and is very productive (Sternberg
et al. 2006). City administrators may also find this
grass helpful in keeping fire ants out of parks, land-
scaped areas, and roadsides. Anderson et al. (2002)
found that water suspensions of perilla (Salvia spp.)
leaves, pine (Pinus spp.) needles, and cedar shav-
Wang et al.: Essential Oils Affect Foraging Behavior of Solenopsis invicta 455
ings can be effective in repelling fire ants. The repel-
lency and toxicity of peppermint oil particles caused
fire ant nests to be abandoned (Appel et al. 2004).
Octanoic acid (Vander Meer et al. 1993), bifenthrin
and tefluthrin (Oi & Williams 1996) were found to
prevent potted plants from becoming contaminated
by the red imported fire ant. Chen (2005) found that
diethyl phthalate and dimethyl phthalate can be ef-
fective in repelling fire ants. Two insect-repellent
terpenoids, callicarpenal and intermedeol, isolated
from the leaves of American beautyberry (Calli-
carpa americana L.; Verbenaceae) and Japanese
beautyberry (Callicarpa japonica Thunb.) were
evaluated (using digging bioassays) and shown to
be effective as repellents against the workers of red
imported fire ants (Chen et al. 2008).
Certain components of the pepper plant Capsi-
cum annuum L. (Solanales: Solanaceae) have long
been known as insect repellents. For example, the
negative effects of extracts from the fruits and seeds
of C. annuum were observed in experiments with
the saw-toothed grain beetle, Oryzaephilus suri-
namensis (L.), (Silvanidae) and the rust-red flour
beetle, Tribolium castaneum (Herbst) (Tenebrioni-
dae), both insect pests of stored products (Iorizzi et
al. 2000). The Himalayan cedar, Cedrus deodara
(Roxb.) G.Don (Pinales: Pinaceae), is a graceful or-
namental evergreen tree that grows extensively on
the slopes of the Himalayas and shows repellent ef-
fects on the nesting of S. invicta. Cedrus deodara
could also supply repellants for use against other
pests. It has been reported that a highly effective oil
obtained from C. deodara caused complete mortal-
ity in Anopheles stephensi (Kumar & Dutta 1987).
In the above-mentioned reports, nest digging be-
havior of workers was used to evaluate the repellent
effects of plant essential oils on fire ants. However,
foraging is one of the most important daily activities
for social insects, including fire ants and other ants.
The repellent effects of plant oils on the foraging be-
havior of the red imported fire ants have not been
tested to date. In this study, the repellent effects of
several plant essential oils on the workers of the
red imported fire ant were evaluated using an ant
foraging bioassay. The results of this study identify
essential oils that significantly deter the foraging
of red imported fire ants. These findings may have
practical significance in quarantine procedures for
fire ants and for public protection.
materials and methods
Materials
Sausage (Xincheng Jinluo Meat Group Co., Ltd.,
Linyi, Shandong, China) which is made mainly
from chicken, starch and sugar was presented as
a lure to observe the foraging behavior and re-
cruitment of S. invicta. The plant essential oils of
Salvia sclarea L.; (Lamiales: Lamiacaeae), C. an-
nuum, Mentha Canadensis L.; (Lamiales: Lamia-
caeae), Mentha longifolia (L.) Huds., C. deodara,
and Pinus spp. were provided by DaMo Chemical
Co. Ltd (Tianjin, China). The essential oil product,
which in Chinese is called Feng You Jing (FYJ)
(Liang Jiefu Pharmaceutical Pte Ltd., Singapore),
was purchased from a Chinese traditional herb
medicine store in the city of Guangzhou, Guang-
dong Province, China in Oct 2010. The percentage
of each compound in FYJ reported on the label was
25% for menthol, 20% for methyl salicylate, 3% for
camphor, 3% for eucalyptus oil, 14% for eugenol
and 35% for liquid paraffin. The hexane and etha-
nol used in the bioassay were Analytical reagent
grade (Sinopharm Chemical Reagent Co. Ltd. and
DaMao Tianjin Reagent Factory).
Field Conditions
The field experiments were performed on the
campus of South China Agricultural University
in Guangzhou, Guangdong, China from Apr to
Jun 2011. The number of inseminated queens
in the polygyne colonies ranged between 2 and
27 and was determined by dissection (Porter
1992). The fire ant density at the study site was
8 mounds/100 m2. On the day of the experiments,
the weather was sunny with air temperature of
25-35 °C and 40~55% RH. The bioassays below
were conducted between 10:00 a.m. to 3:00 p.m.
Deterrent Effects of Essential Oils on the Food-
Searching Behavior of the Red Imported Fire Ant
Based on the report of Chen et al (2008) on the
bioassay of repellency against the digging behavior
of fire ant workers and on our preliminary experi-
ments, we diluted essential oils with hexane to 10-6
mL/mL for the following bioassay. First, 100 μL of
oil solution or hexane (control) was dropped onto the
center of a 70 mm-diam piece of filter paper using a
pipette. Five s later, 8 randomized filter paper piec-
es were each placed 15 cm from the nest border. A
0.5 g piece of sausage was then placed on each piece
of filter paper (Fig. 1). The search time (the time
required for one of the sausages to be discovered)
and the number of recruited workers of the fire ants
for each food source were recorded. A video camera
(Nikon D90 system) was positioned above each
mound by a tripod in order to record the numbers of
workers on the filter papers at different times. Each
treatment was repeated with 5 nests.
Interference Effects of Essential Oils on Food Transport
by Red Imported Fire Ants
Prior to the test, the sausages were cut into slic-
es weighing 0.5 g and then divided into 8 smaller
fragments each ~ 62.5 mg. A total of 100 μL of oil
solution or hexane (control) was dropped onto the
center of a piece of filter paper using a pipette. Five
456 Florida Entomologist 97(2) June 2014
s later each of the filter paper piece was randomly
placed on 1 of the 8 locations (Fig. 1) each exactly
15 cm from the nest perimeter. Eight fragments of
sausage (each approximately 0.625 mg) were then
placed on each piece of filter paper. We recorded the
time and observed and counted the number of sau-
sage fragments transported by the workers within
consecutive 5 min intervals until all of the sausage
pieces had been completely removed. Each treat-
ment was repeated with 5 nests.
Statistical Analysis
The differences in the search times, the number
of recruits and number of sausage pieces transport-
ed under the different treatments, exposure times,
and at different fire ant nests were measured. All
data were tested for normal distribution by the Sha-
piro-Wilk test and for homogeneity of variances by
Levene’s test. Two-way analysis of variance (ANO-
VA) was performed to compare the search time of
workers and the time required for food transporta-
tion exposed to different essential oils. Three-way
ANOVA was used to compare the number of fire ant
workers recruited and the amounts of food trans-
ported from different essential oil treated papers at
different exposure times and different nests. When
necessary, one way ANOVA was performed for each
single factor. All of the statistical analyses were per-
formed using the SPSS 13.0 software package.
results
Search Times with Different Essential Oils
The experiment showed that the foraging be-
havior of the fire ants was affected by the 7 types
of essential oils (Fig. 2). All of the essential oils
Fig. 1 Schematic map for placing baits on essential
oil-treated pieces of filter paper each equidistant from
the fire ant nest in the field bioassay of the repellent
effects of various essential oils. A dark circle represents
the fragment of sausage on a piece of filter paper (white
circle). The distance from the edge of each filter paper
piece to the nest was 15 cm.
Fig. 2. Time (min) required for Solenopsis invicta workers to discover a 0.5 g fragment of sausage placed on a
piece of filter paper treated with an essential oil. Each piece of filter paper was placed 15 cm from the perimeter of
the fire ant mound. FYJ is a mixture of menthol, methyl salicylate, camphor, eucalyptus oil and eugenol. Means (±
SE) followed by the same letter are not significantly different (LSD) at level of 0.05.
Wang et al.: Essential Oils Affect Foraging Behavior of Solenopsis invicta 457
except that of Mentha longifolia significantly pro-
longed the time required for the workers to arrive
at the sausage compared to the untreated control.
However only C. annuum significantly prolonged
the search time more than M. longifolia, and the
prolongation of the search time of C. annuum es-
sential oil was not significantly greater than those
of the essential oils of C. deodara, FJY, Pinus
spp., S. sclarea, or M. canadensis. In addition, we
observed that the fire ants made repeated visits
to inspect the sausage. This behavior seemed to
affect the subsequent recruitment.
The results showed that the search times of
workers for the sausage were affected significantly
by both the fire ant nest (F = 21.487, df = 4, 28; P
< 0.0001) and essential oil type (F = 10.382, df = 7,
28; P < 0.0001). Search times for the fire ants with
the C. annuum oil and C. deodara oil treatments
were 3.64 min and 2.98 min, respectively; these
values were greater than the control (1.53 min).
Recruitment of Fire Ant Workers in Different Essential
Oil Treatments
The numbers of worker ants recruited (Table
1) during the observation periods differed signifi-
cantly with different essential oils (F = 49.565, df
= 7, 280; P < 0.0001), exposure time (F = 139.701,
df = 6,280; P < 0.0001) and the fire ant nests (F =
30.540, df = 4, 280; P < 0.0001). No significant dif-
ferences in recruitment occurred during the initial
10 min of exposure time. When the exposure time
exceeded 10 min, the number of workers at the
sausage fragments differed significantly among
the different essential oil treatments. Capsicum
annuum oil had the greatest effect, followed by
the C. deodara oil, and S. sclarea oil. In the pep-
permint treatment, the number of worker ants
recruited was significantly less than the control
when the exposure time is greater than 10 min. In
the treatments of C. deodara oil and S. sclarea oil
treatments, the numbers of worker ants recruit-
ed were significantly less than in the control for
exposure times greater than 15 min and 20 min,
respectively. Worker recruitment was significantly
lower when exposure times were greater than 25
min and 35 min for the Pinus spp. oil and FYJ
treatments, respectively. These results show that
recruitment after discovery of food depend on both
the exposure time and the type of oil used. In our
test, the C. annuum oil and C. deodara oil showed
better recruitment deterrence effects on the work-
ers than the other essential oils tested (Table 1).
The Amount of Food Transported and the Time Required
for Food Transportation under Different Essential Oil
Treatments
The amount of food transported by the worker
ants (Table 2) showed an increasing trend with
table 1. the numbers of SolenopSiS invicta Workers attracted to sausage fragments on 70 mm-diam pieces of filter paper each treated With a standard
amount of an essential oil and placed 15-cm from the nest. the number of recruits Was recorded in 5 min intervals from the time of placement.
Plant oils
Number of Workers
5 min 10 min 15 min 20 min 25 min 30 min 35 min
Capsicum annuum 2.00 ± 0.87 aA 2.60 ± 1.47 aA 5.60 ± 2.48 aAB 6.80 ± 2.62 aAB 9.00 ± 4.24 aAB 10.60 ± 4.81 aB 13.00 ± 6.12 aB
FYJ 3.80 ± 2.02 aA 7.40 ± 4.14 abA 14.20 ± 4.75 abAB 20.60 ± 6.03 bcBC 28.20 ± 7.25 cdCD 33.40 ± 7.58 bcD 41.40 ± 7.97 bD
Mentha longifolia 3.40 ± 1.18 aA 9.20 ± 4.10 abA 20.40 ± 6.53 cB 30.80 ± 7.32 cC 42.00 ± 5.07 eD 51.20 ± 4.37 dE 62.00 ± 3.28 cF
Mentha canadensis 3.80 ± 0.92 aA 11.00 ± 4.18 bAB 19.40 ± 7.06 bcBC 27.80 ± 7.41 cCD 36.60 ± 8.51 deDE 45.60 ± 8.22 cdEF 56.00 ±7 .23 cF
Salvia sclarea 2.60 ± 1.47 aA 4.40 ± 2.48 abA 9.00 ± 6.24 abAB 12.60 ± 8.54 abAB 19.00 ± 8.63 abBC 29.20 ± 10.25 bCD 37.80 ± 8.10 bD
Cedrus deodara 2.20 ± 0.92 aA 4.40 ± 2.04 abA 7.20 ± 3.06 aAB 11.80 ± 3.98 abBC 13.80 ± 4.56 abBCD 16.60 ± 4.57 aCD 20.20 ± 5.46 aD
Pinus spp. 3.40 ± 1.28 aA 8.60 ± 3.15 abA 12.80 ± 5.32 abA 18.80 ± 4.26 bcB 24.20 ± 3.95 bcB 31.40 ± 2.94 bC 39.60 ± 2.77 bD
Control 3.60 ± 1.18 aA 9.60 ± 3.07 bAB 18.20 ± 4.51 bcB 28.40 ± 5.84 cC 37.80 ± 6.21 deD 45.20 ± 5.95 cdD 56.40 ± 5.82 cE
*Means in the same column followed by the same small letter or in the same row followed by the same capital letter are not significantly different (LSD) at level of 0.05.
458 Florida Entomologist 97(2) June 2014
the elapse of time (F = 5.147, df = 6, 280; P <
0.0001) and differed significantly for the same es-
sential oil treatment (F = 28.098, df = 7, 280; P <
0.0001) and the fire ant nest (F = 154.085, df = 4,
280; P < 0.0001). No significant differences were
found in the amounts of food transported for ex-
posure times of 5 min and 10 min. For exposure
times greater than 10 min, the amounts of food
transported differed significantly in the different
essential oil treatments (Table 2). Cedrus deoda-
ra oil had the greatest suppressive effect on the
amount of food transported by the workers fol-
lowed by the C. annuum oil. When the exposure
time was 10 min or less, no food transport was
found in the experiments using C. deodara oil and
C. annuum oil; however, when the time interval
was as long as 35 min, the average amount of food
transported was 4.8 and 5.4 pieces, respectively.
In contrast, the amount of food transported was 7
pieces or more for the other essential oils tested.
The time required to transport all of the food
differed (Fig. 3) markedly among the different
essential oil treatments (F = 15.257, df = 7, 28;
P < 0.0001) and the fire ant nest (F = 2.774, df
= 7,28; P = 0.046). It should be noted that all of
the treatments were presented in the same habi-
tat. The time required to transport all the food
was the greatest under the C. annuum oil treat-
ment (53.87 min), followed in declining order by
the C. deodara oil (50.23 min), the M. canadensis
oil (41.98 min), FYJ (40.57 min) and the control
(27.22 min). Thus, the C. annuum oil and the
C. deodara oil have significant impacts on the
amount of food transported and the time required
to transport all of the food.
discussion
Foraging trails of S. invicta colonies range
in length from 1 cm to 2 m (Markin et al. 1975),
which indicate the distance from the food re-
source to the tunnel opening. Therefore, we set up
the sausage only 15 cm away from the nest border
to achieve a uniform the trail length. Our field ob-
servations confirmed that the foraging trails were
almost between 5 cm to 10 cm in our experiments.
During foraging, fire ant workers detect the smell
of food with their antennae, and the type, location
and weight of the food can affect their subsequent
recruitment behavior (Xu et al. 2007). The results
of the study showed that the presence of essential
oils reduced the number of ants arriving at the
filter paper, which in turn, influenced the search
time, the recruitment and the transport of the
food by the workers.
The essential oils were clearly repellent
against the foraging of fire ants, and the type of
essential oil used made a substantial difference
in this effect. Under outdoor conditions, the C.
annuum oil and C. deodara oil were the most
effective repellents among the 7 essential oils
table 2. the numbers of sausage fragments transported bY SolenopSiS invicta Workers from 70 mm-diam pieces of filter paper each treated With a stan-
dard amount of an essential oil and placed 15-cm from the nest. the number of sausage fragments removed from each piece of filter paper
Was recorded in 5 min intervals from time of placement.
Plant oils
Sausage slices transported
5 min 10 min 15 min 20 min 25 min 30 min 35 min
Capsicum annuum 0.00 ± 0.00 aA 0.00 ± 0.00 aA 0.60 ± 0.39 aA 1.40 ± 0.63 abAB 2.80 ± 1.16 abBC 4.00 ± 1.22 abCD 5.40 ± 1.07 aD
Cedrus deodara 0.00 ± 0.00 aA 0.00 ± 0.00 aA 0.80 ± 0.59 abAB 1.00 ± 0.71 aAB 2.00 ± 1.11 aBC 3.60 ± 1.37 aCD 4.80 ± 1.16 aD
Mentha longifolia 0.60 ± 0.39 abA 2.20 ± 0.92 bB 4.60 ± 1.07 cC 6.20 ± 0.92 deD 7.40 ± 0.38 eD 9.00 ± 0.50 dE 10.00 ± 0.00 cE
Salvia sclarea 0.00 ± 0.00 aA 1.00 ± 0.5 abA 2.20 ± 0.77 abcAB 3.40 ± 1.07 abcBC 4.60 ± 1.28 bcCD 6.20 ± 1.45 bcDE 7.40 ± 1.91 abE
FYJ 0.40 ± 0.63 abA 2.00 ± 1.66 bAB 4.00 ± 2.18 cABC 5.60 ± 2.16 cdeBCD 7.00 ± 1.87 cdCD 8.50 ± 1.35 cdD 7.50 ± 0.71 abCD
Pinus spp. 0.20 ± 0.32 abA 0.80 ± 0.59 abA 2.20 ± 1.05 abcAB 3.80 ± 1.61 bcdBC 5.60 ± 1.70 cdCD 7.00 ± 1.65 cdD 7.50 ± 1.91 abD
Mentha canadensis 1.00 ± 0.71 bA 2.00 ± 1.33 bA 3.0 0± 1.41 abcAB 5.00 ± 1.11 cdeBC 7.00 ± 1.22 cdCD 8.00 ± 1.22 cdD 9.00 ± 1.58 cD
Control 0.40 ± 0.39 abA 1.20 ± 0.92 abAB 3.20 ± 1.16 bcB 6.60 ± 1.63 eC 7.25 ± 0.68 cC 9.50 ± 0.41 dD 10.00 ± 0.00 cD
*Means in the same column followed by the same small letter or in the same row followed by the same capital letter are not significantly different (LSD) at level of 0.05.
Wang et al.: Essential Oils Affect Foraging Behavior of Solenopsis invicta 459
tested. The concentration of essential oils used
in this experiment was 10-6 ml/ml, but different
concentrations may alter the effectiveness of a
repellent (Chen 2009). Future research should
consider this possibility. Our study also revealed
that the number of worker ants involved in forag-
ing activities tended to increase over time. This
may due to the gradual decrease of the repellent
effect of essential oils. It is probable that lower
repellency occurred because the outdoor air-flow
accelerated the evaporation of the essential oils.
Perhaps micro-capsules could be developed to en-
sure a greater stability of the repellent effects.
Plants use secondary metabolites to resist
potential damage by herbivores. These second-
ary metabolites are the result of the long-term
coevolution between herbivorous animals and
plants (Hartmann 2004). Compared with chemi-
cal pesticides, plant essential oils can serve as
environmentally friendly pesticides because they
are usually safer and less toxic to humans and
domestic animals, and they are more readily de-
graded in the environment (Isman 2000). Many
scientists are beginning to focus on essential oils
to develop efficient new, minimally toxic pes-
ticides for use against pests that threaten both
health and agricultural production. According to
Chen (2009), FYJ could be improved to act as a
repellent against red imported fire ants workers.
Our results indicated that the repellent proper-
ties (especially the persistence effect) of C. an-
nuum and C. deodara were stronger than that
of FYJ. Although C. annuum and C. deodara
showed effectiveness in repelling foraging work-
ers, the number of workers recruited was greater
after 30 min than 10 min. Modification of C. ann-
uum and C. deodara oils to produce longer effects
over longer periods are needed if these plant oils
are to be used as repellents for preventing food
contamination by fire ants. These oils, may also
prove useful in repelling fire ants from nesting
soil (Chen & Allen 2006).
acknoWledgments
We thank Rensen Zeng at the South China Agri-
cultural University for constructive comments regard-
ing this manuscript. The current study was supported
by the National Basic Research Program of China
(2009CB119200).
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