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Journal of Economic Entomology, 2017, 18

doi: 10.1093/jee/tox107
Research article
Household and Structural Insects

Development of Liquid Bait With Unique Bait Station for


Control of Dolichoderus thoracicus (Hymenoptera:
Formicidae)
Chung-Chi Lin,1 Tsai-Wei Chang, Hung-Wei Chen, Chung-Han Shih, and Po-Cheng Hsu
Department of Biology, National Changhua University of Education, No. 1, Jin-De Rd., Changhua City, 50007, Taiwan (cclin@
cc.ncue.edu.tw; sj801112@yahoo.com.tw; mark77529@livemail.tw; harry820902@gmail.com; s810093125@gmail.com), and
1
Corresponding author, e-mail: cclin@cc.ncue.edu.tw

Subject Editor: Arthur Appel


Received 12 December 2016; Editorial decision 20 March 2017

Abstract
The Dolichoderus thoracicus (Smith) has seriously infested urban, village, and agricultural environments in
Taiwan. To develop optimal bait for the effective control of D. thoracicus, we investigated the feeding prefer-
ences of this ant at different sugar and amino acid concentrations in a laboratory setting. The number of feeding
workers was significantly higher for 20% sucrose compared with that for other sugars at 10% and 20% concen-
trations. Furthermore, among all tested concentrations of monosodium glutamate (MSG; 1%, 3%, 5%, and
10%), the highest number of ants were attracted by 1% MSG with 20% sucrose. The consumption of each bait
solution was also measured, and the results were similar to those for ant number. Regarding toxicants, the tox-
icity and repellency of 3% boric acid and borax bait were evaluated individually. Both 3% boric acid and borax
demonstrated no repellency against D. thoracicus workers. The LT50 of the 3% boric acid bait was 4.87 d,
whereas that of 3% borax bait was only 1.56 d. Thus, 3% boric acid showed further delayed action, facilitating
workers spreading the toxicant to their nestmates through trophallaxis. In combination with our unique bait sta-
tion, we propose an optimal bait formulation comprising 20% sucrose, 1% MSG, and 3% boric acid as a promis-
ing candidate for controlling D. thoracicus.

Key words: Dolichoderus thoracicus, bait, boric acid, feeding preference, pest ant

The Dolichoderus thoracicus (Smith) (Hymenoptera: Formicidae) main methods used for reducing ant infestations, can rapidly reduce
was described by Smith (1860), and is native to southeast Asia. It the number of pest ants; however, this method has several limita-
opportunistically nests in folded leaves or leaf litter on the ground tions: 1) Their use only causes partial suppression because they
(Khoo and Chung 1989). This ant has been evaluated as a biocon- mostly kill workers which foraged outside; the queen, workers, and
trol agent to suppress populations of pests in economic crops, such larvae in the nest remain unaffected (Knight and Rust 1990). 2) To
as cocoa and sapodilla (Khoo and Ho 1992, Van Mele and Cuc maintain their efficacy, sprays must be reapplied, making this
2001). This species has also been deliberately introduced into method time- and cost-intensive (Rust et al. 1996). 3) These sprays
Malaysia, Argentina, and Hawaii (Taylor 1987, Reimer 1994, are also toxic to beneficial, nontarget organisms; moreover, the
Nishida 2002). In Taiwan, D. thoracicus is mainly distributed in chemical residues cause environmental pollution and are retained in
bamboo forests, and gardens. It can also invade indoor and outdoor crops, indirectly damaging human health (Smith et al. 1996). Thus,
areas associated with human activity by establishing new spreading an increasing number of studies have focused on the use of toxic
and foraging routes in water pipes and wires. These manmade areas baits as an alternative method for controlling pest ants (Lee and Lee
provide this pest suitable habitats, like the edges of windows, ceil- 2002, Daane et al. 2008).
ings, and beams. Besides, D. thoracicus can obtain sugar sources Toxic baits are composed of a delayed-action insecticide incor-
from flowers and hemipteran insects in manmade gardens or yards. porated into a palatable food. By using the characteristics of recruit-
To date, this pest severely reduces the quality of human life in ment and trophallaxis in ants, baits target the entire ant colony
Taiwan. However, no effective method of control has been rather than just foragers (Silverman and Tai 2001). In addition, ra-
reported. ther than being sprayed widely, a small quantity of the insecticide is
The most common strategy for managing pest ant populations is placed in the bait station and delivered from a point source, thus
the chemical control. Treatment with residual sprays, one of the reducing environmental damage (Hooper-Bui and Rust 2000).

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2 Journal of Economic Entomology, 2017, Vol. 0, No. 0

Knowledge of the precise location of ant nests is not required when


baits are deployed (Chong and Lee 2009).
Effective ant baits generally have these characteristics: ant-
preferred food ingredients, a slow-acting insecticide, and nonrepel-
lent preservatives (Cherrett and Lewis 1974, Rust et al. 2004,
Silverman and Brightwell 2008). Baits are usually formulated as li-
quids, granules, and gels (Silverman and Tai 2001, Hara et al.
2014). To date, many studies have focused on the optimal concen-
tration of toxicants to effectively control pest ants (Hooper-Bui and
Rust 2000, Ulloa-Chac! on and Jaramillo 2003). Other studies have
investigated the various bait matrices impacting feeding preferences
of ants (Hara et al. 2014). However, few studies have focused on
food source preferences and food concentrations for D. thoracicus.
The ingredients of ant baits are essentially carbohydrates and
proteins. Several studies have shown that these two components
strongly affect feeding preference in ants (Ricks and Vinson 1970,
Kaspari and Yanoviak 2001). Various ant species may specifically
sense and forage for their preferred carbohydrates (Vander Meer
et al. 1995). In addition, Hahn and Wheeler (2002) demonstrated
that arboreal ants preferentially select protein-based baits, whereas Fig. 1. Schematic of the liquid bait station developed in our laboratory.
terrestrial ants tend to feed on carbohydrate-based baits. Therefore,
an objective of our research was to analyze the degree of attraction
of D. thoracicus to baits of various formulations. In terms of the Taiwan) as nitrogen source, and 5-ml water vial plugged with cotton
slow-acting insecticides, boron compounds (boric acid or borax) are as water source. These foods and water were supplied every day.
toxicants that have been used for controlling ants for over a century Ant colonies were maintained in the laboratory (at about 25 ! C and
(Riley 1889). Boron compound-based baits have been assessed for a photoperiod of 12:12 [L:D] h) for subsequent experiments.
their toxic effects on many ant species, such as carpenter ants,
Camponotus abdominalis (L.) and Argentine ants, Linepithema Bait Station
humile (Mayr) (Klotz et al. 1996, 1997). Although the toxicity of We used a liquid bait station developed in our laboratory. It was
borate-containing baits toward several ant species has been well- composed of two parts: an attractant platform and a 50-ml centri-
studied, little is known about their effects on D. thoracicus. Because fuge tube to hold the bait solution (The bait solutions which we
asymmetrical feeding preference toward two baits with boric acid used in this study are described in the following subsection). A small
and borax may depend on the ant species (Sola et al. 2013), the se- hole was drilled in the lid of the centrifuge tube before use. The
cond objective of this study was to determine the toxicity of boric drilled end of the tube is attached to the notch (Fig. 1, the upper
acid- and borax-based baits on D. thoracicus. half) of upside-down bait station. Then, both the centrifuge tube
In the wild, D. thoracicus favor honeydew or extrafloral necta- and bait station were inverted together to allow the solution to fill
ries, which are liquid food sources (Ho and Khoo 1997, Way and up the groove (Fig. 1, the lower half) when we deployed the baits.
Bolton 1997). Thus, we selected a liquid form, rather than granule This bait station can continue providing stable liquid bait solution
form, of bait for our study. We investigated the feeding preference for up to 7 d under general conditions. The groove was designed to
of D. thoracicus toward liquid baits with different sugar sources and allow a large number of ants to feed on the liquid bait simultan-
concentrations, additional nitrogen nutrients, and a preservative eously (Fig. 2A, B).
under laboratory condition. And the lethal effect (toxicity) of the
toxicants was evaluated. This is the first study to facilitate the devel- Bait Preference and Repellency Tests
opment of sustainable tools for managing D. thoracicus outbreaks Tests were conducted to determine the feeding preference by D.
in urban and agricultural environments. thoracicus to bait solutions: 1) Sucrose, fructose, and maltose were
individually dissolved in reverse osmotic water (RO water, which
was purified by passing through semipermeable membrane to re-
Materials and Methods
move ions and large particles) to prepare 10% and 20% (mass/vol-
Insects ume) sugar solutions. 2) For the nitrogen source (amino acid)
The bamboo branches or culms, where D. thoracicus lived, were cut preference, solutions containing 1%, 3%, 5%, and 10% (mass/vol-
and placed into a zip-lock bag in Shihu Township, Miaoli County, ume) monosodium glutamate (MSG) with 20% sucrose, and control
Taiwan. The branches and culms (wild nests) were transferred to the (20% sucrose) solution were prepared as additional nitrogen treat-
Social Insect Laboratory at National Changhua University of ments. 3) To prevent microbial contamination and prolong liquid
Education, Taiwan. Ants from different nests were placed separately solution life span (Buczkowski et al. 2014), we compared the bait
in large plastic boxes (47 by 34 by 26 cm); the upper part of the solution (20% sucrose and 5% MSG) containing 0.3% (mass/vol-
inner walls was smeared with Fluon (polytetrafluoroethylene; ume) benzoic acid as a preservative with a control solution (20% su-
Northern Products, Inc., Woonsocket, RI) to prevent the ants from crose 5% MSG). This test was designed to determine whether
escaping. Bamboo branches or culms were set to provide space for benzoic acid was repellent to D. thoracicus. 4) Crystalline boric acid
nesting. Colonies were provisioned with plastic dishes containing (H3BO3) and borax (Na2B4O7) were dissolved in 20% sucroseRO
cotton which was wet with 20% sucrose solution as carbon source, water with 1% MSG to a 3% (mass/volume) concentration each. To
pieces of egg pudding (Uni-President Enterprise Co., Tainan, verify that the active ingredients were not repellent, we also
Journal of Economic Entomology, 2017, Vol. 0, No. 0 3

Fig. 2. Images of D. thoracicus workers feeding on liquid baits containing (A) 20% sucrose and (B) 20% sucrose and 1% MSG.

compared these two toxic baits with a control solution (20% dead ants was recorded daily to further calculate the survival rate:
sucroseRO water with 1% MSG). Each solution was added to the [(500 # the number of dead ants)/500 $ 100%].
50-ml centrifuge tube before installation into the bait station.
A total of 600 ant workers were haphazardly selected and placed Statistical Analysis
in plastic boxes containing bamboo branches and RO water in each The results of the feeding preference tests for sugar and amino acid
test colony. The ants were allowed to acclimatize to the nest for 1 sources were analyzed using one-way analysis of variance
wk and were then starved for 1 d before tests were initiated. The dif- (ANOVA), followed by the BonferroniDunn post hoc test, whereas
ferent bait solutions were then individually introduced into the bait the results of tests for benzoic acid as a preservative were analyzed
stations and separately deployed in different test colonies. The ants using Students t test. To calculate the LT50 of the two boron-based
were allowed to feed ad libitum. Feeding activity was monitored by compounds for workers, data were corrected using the formula de-
time-lapse camera (TLC200, Brinno, Taipei, Taiwan), and images veloped by Abbott (1925) and analyzed using probit analysis
captured every 30 s for 1 h. The average number of feeding ants was (Raymond 1985). The survival analyses for different treatments
evaluated from 20 images per 10 min; only the ants that stayed on were computed using the KaplanMeier estimator (Kaplan and
the bait station and touched the bait solutions with its mouthpart Meier 1958), and the resulting survival curves were plotted. These
were counted. The six figures acquired from the 10-min intervals curves were further compared and any obvious differences were
during 1 h represented the total number of feeding ants. Six repli- examined using the log-rank test (Cox and Oakes 1984), where
cates were performed for each treatment. P < 0.05 was considered statistically significant.

Bait Consumption
Bait consumption was determined by visual observation of the
Results
change in the scale on the centrifuge tube. The drop distance of li- Bait Acceptance
quid surface was measured by Vernier calipers and converted into 20% sucrose solution was more attractive than any other sugar baits
the volume (6 0.1 ml). The mean consumption of each solution for (Fig. 3; F 6.86, df 2, 39; P < 0.01): 435.3 6 25.2 workers con-
1 d was calculated from six replicates. Water lost through evapor- sumed the bait with 20% sucrose for 1 h. Sucrose was more attractive
ation was subtracted from the volume in the centrifuge tube as a cor- than the other two sugar foods. Bait consumption showed a similar
rection factor. The evaporative water loss was detected by pattern (Fig. 4; F 4.68, df 2, 39; P < 0.05): consumption was
depositing the bait station in adjacent nest boxes without ants and highest in the 20% sucrose solution (18.8 6 1.2 ml/d), followed by
measuring the volumetric reduction as described above. the 10% sucrose solution (13.6 6 1.7 ml/d). For other sugar solutions,
the consumption was lower than 8 ml. 20% sucrose was the most pre-
Toxicity Tests ferred by D. thoracicus and was selected for the subsequent tests.
Two toxic baits (3% boric acid or borax in 20% sucroseRO water
with 1% MSG) were separately placed in test ant colonies. Each test Attractiveness of Baits
ant colony placed 500 workers and acclimatization process for ants Figure 5 illustrated the number of foraging D. thoracicus at different
was identical to that for the feeding preference tests. After starvation MSG concentrations in the 20% sucrose solution. The solution con-
for 1 d, the dead ants were removed with new workers before the taining 1% MSG with 20% sucrose attracted a significantly greater
tests were initiated. Each bait or control (20% sucroseRO water number of ants compared with the control (F 22.08, df 4, 25;
with 1% MSG) was provided to the ants for 8 d. The number of P < 0.001): this solution attracted 525.3 6 27.8 ants, indicating an
4 Journal of Economic Entomology, 2017, Vol. 0, No. 0

Fig. 3. Numbers of D. thoracicus workers (mean 6 SE) feeding on liquid baits each containing 10% and 20% sucrose, fructose, and maltose per hour. RO water
was used as control. Means followed by the same letter indicate nonsignificant difference (one-way ANOVA, BonferroniDunn post hoc, F 6.86; df 2, 39;
P < 0.01; n 6).

Fig. 4. Daily consumption (mean 6 SE) of bait solutions each containing 10% and 20% sucrose, fructose, and maltose. RO water was used as control. Means fol-
lowed by the same letter indicate nonsignificant difference (one-way ANOVA, BonferroniDunn post hoc, F 4.68; df 2, 39; P < 0.05; n 6).

&1.25-fold increase compared with the control (420.8 6 18.7 ants). solution containing 1% MSG in 20% sucrose was selected as the
At higher concentrations of MSG (3%, 5%, and 10%), the number optimal food ingredient formulation for a bait solution to attract
of feeding ants did not increase significantly compared with the con- D. thoracicus in this study.
trol. In addition, bait consumption for 1 d for these treatments and
the controls demonstrated a similar trend (Fig. 6; F 29.3; df 4,
25; P < 0.001): the 1% MSG solution showed the highest daily con- Repellency of Baits With Benzoic Acid
sumption (26.9 6 1.9 ml/d, significantly differing from that of the To assess whether benzoic acid was repellent to D. thoracicus, we
control [18.1 6 1.7 ml/d]), whereas the consumption of other MSG added 0.3% benzoic acid to 20% sucrose containing 1% MSG and
solutions was similar to that of the control. Taken together, the performed the preference tests in comparison to a control (1% MSG
Journal of Economic Entomology, 2017, Vol. 0, No. 0 5

Fig. 5. Numbers of D. thoracicus workers (mean 6 SE) feeding on solutions containing 20% sucrose each with 1%, 3%, 5%, and 10% MSG per hour. Solution con-
taining 20% sucrose was used as control. Means followed by the same letter indicate nonsignificant difference (one-way ANOVA, BonferroniDunn post hoc,
F 22.08; df 4, 25; P < 0.001; n 6).

Fig. 6. Daily consumption (mean 6 SE) of bait solutions each 20% sucrose each with 1%, 3%, 5%, and 10% MSG. Solution containing 20% sucrose was used as
control. Means followed by the same letter indicate nonsignificant difference (one-way ANOVA, BonferroniDunn post hoc, F 29.3; df 4, 25; P < 0.001; n 6).

in 20% sucrose solution). The results (Fig. 7) revealed that the num- to D. thoracicus: the results (Fig. 8) revealed that the numbers of
bers of ants feeding per hour on bait solutions with and without ben- ants feeding on the treatment and control bait solutions per hour did
zoic acid were similar (t 1.6, df 10; P 0.14), indicating 0.3% not differ significantly (F 2.21, df 2, 15; P 0.144). We next
benzoic acid was not affecting attractant to D. thoracicus. evaluated the toxic effects of borax and boric acid on D. thoracicus.
Table 1 presents the survival rates of workers in baits containing
3% borax, 3% boric acid, and the control every day for 8 d. The
Repellency and Toxicity of Boron Baits toxicity of 3% borax was higher than that of the 3% boric acid or
First, we verified that our optimal bait solutions (1% MSG in 20% control during the test. The LT50 of 3% borax was 1.56 d, three
sucrose) with 3% borax or boric acid were not affecting attractant times shorter than that of boric acid (4.87 d). According to the
6 Journal of Economic Entomology, 2017, Vol. 0, No. 0

Table 1. Survival rate of D. thoracicus workers fed solutions con-


taining 20% sucrose, 1% MSG, and 3% borax or boric acid for 8 d

Days Borax (3% w/v) Boric acid (3% w/v) Controla


0 100% 100% 100%

1 70.7% 6 22.0% 98.8% 6 0.7% 98.8% 6 1.2%


2 26.3% 6 9.8% 96.1% 6 2.9% 96.2% 6 2.2%
3 6.6% 6 2.5% 85.8% 612.7% 93.8% 6 4.2%
4 4.2% 6 1.9% 67.2% 6 9.2% 92.4% 6 1.0%
5 3.4% 6 2.3% 42.1% 6 1.2% 90.2% 6 1.6%
6 2.4% 6 2.0% 27.0% 6 4.8% 89.2% 6 1.1%
7 0.0% 8.8% 6 2.9% 88.2% 6 2.6%
8 0.0% 1.3% 6 0.6% 87.6% 6 1.4%

a
Control: Solutions containing 20% sucrose and 1% MSG.

most attractive sugar, followed by glucose. The variations in sugar


preference may be associated with the availability of native food re-
sources in ant habitats. The major liquid food source for ants is nec-
Fig. 7. Numbers of D. thoracicus workers (mean 6 SE) feeding on liquid baits tar from plants or honeydew from mealybugs, both of which
containing 20% sucrose and 1% MSG with or without benzoic acid. Solution contain different compositions and concentrations of sugar and
containing 20% sucrose and 1% MSG was used as control (t test, t 1.6; amino acids depending on the species (Fischer and Shingleton 2001,
df 10; P 0.14; n 6).
Bluthgen et al. 2004). Considering amino acids, D. thoracicus pre-
ferred 1% MSG in 20% sucrose; however, no feeding preference
was noted for the higher concentrations of MSG (3%, 5%, and
10%) in 20% sucrose compared with the control. By contrast, in a
previous study, Solenopsis geminata (F.) preferred amino acid-rich
nectar mimic over a solution with a lower amino acid content
(Lanza et al. 1993). Interspecific differences have been noted in the
feeding preferences for amino acids among some ant species; for in-
stance, Oecophylla smaragdina (F.) was found to favor arginine
over glutamic acid and Pheidole platypus Crawley preferred sugar
only over sugar with serine (Bluthgen and Fiedler 2004). Notably,
amino acids have demonstrated repellency against ants in some
cases; therefore, each ant species can be concluded to have specific
amino acid preferences. Dolichoderus thoracicus may favor glu-
tamic acid, dissociated from the MSG provided as the nitrogen
source. Furthermore, ants dietary choices are affected by colony de-
velopment and requirements (Edwards and Abraham 1990,
Dussutour and Simpson 2008). Larvae prefer amino acids for their
growth, whereas workers require carbohydrates as metabolic fuel
for their activity and competitive performance (Cassill and
Fig. 8. Numbers of D. thoracicus workers (mean 6 SE) feeding on bait solu- Tschinkel 1999, Grover et al. 2007). Here, the consumption of 3%,
tions containing 20% sucrose, 1% MSG, and 3% boric acid or borax. Solution
5%, and 10% MSG did not differ significantly compared with the
containing 20% sucrose and 1% MSG was used as control (one-way ANOVA,
BonferroniDunn post hoc, F 2.21; df 2, 15; P 0.144; n 6). control, probably because 1% MSG sufficiently fulfilled the amino
acid requirement of D. thoracicus. Taken together, both physio-
logical demands and resource availability contributed to the feeding
results of the KaplanMeier estimator and the log-rank test, the sur-
preferences of the ants. Furthermore, adding 1% MSG to a 20% su-
vival curves of both borax and boric acid were significantly lower
crose solution strengthened the attraction of D. thoracicus. This
than that of the control (P < 0.05; Fig. 9). Approximately 50% of
phenomenon was also observed in Camponotus japonicus Mayr,
the ants feeding on the 3% borax bait solution died at 1.5 d; this
where glycine had a synergistic role in the gustatory response to glu-
duration may be insufficient for ant foragers to bring food to their
cose (Wada et al. 2001).
nest for trophallaxis. Thus, we suggest the addition of 3% boric acid
Boron-based compounds have widely been used in baits to con-
in a solution comprising 20% sucrose and 1% MSG to create an ex-
trol pests because they are water soluble, have delayed action, and
cellent bait solution for controlling D. thoracicus.
are less toxic to mammals than other insecticides (Quarles 1992,
Klotz and Williams 1996). They disrupt the regulation of water bal-
ance in ants, causing them to consume more bait to contend with de-
Discussion hydration (Klotz and Moss 1996). The gradual accumulation of the
Carbohydrates attract ants in species-dependent manners (Vander toxic effect induces cell death in the midgut, postpharyngeal glands,
Meer et al. 1995). In this study, D. thoracicus favored sucrose liquid and Malpighian tubules (Sumida et al. 2010). Here, a 90% reduc-
bait. This result is consistent with that of other ant species. Bluthgen tion was observed in the number of D. thoracicus workers treated
and Fiedler (2004) reported that for 23 ant species, sucrose was the with a 3% boric acid bait solution for 1 wk; the same mortality was
Journal of Economic Entomology, 2017, Vol. 0, No. 0 7

Fig. 9. Survival curves of ants treated with bait solutions containing 20% sucrose, 1% MSG, and 3% borax or boric acid and with control (20% sucrose and 1%
MSG) for 8 d (KaplanMeier estimate method, log-rank test, P < 0.05; n 6).

achieved within only 3 d with a 3% borax bait solution. To elimin- At present, D. thoracicus is mainly distributed in southern Asia
ate an entire ant colony by using a bait solution, sufficient time must and has increasingly infested human environments in Taiwan. These
be allocated for workers to transfer the toxicant to other nest mem- ants have also been discovered at ports of entry in New Zealand
bers through trophallaxis (Rust et al. 2004). We observed that 3% (Ward et al. 2006). Way and Bolton (1997) reported that the
boric acid has more delayed action than does 3% borax. D. thoracicus population increased excessively after it became the
The toxicity of boric acid to ants has been evaluated in many sole local species in Malaysian coconut trees. D. thoracicus may
studies. Ulloa-Chac! on and Jaramillo (2003) reported that all work- soon be considered an invasive species. Our study provided useful
ers, broods, and queens of the Tapinoma melanocephalum (F.), died information for developing an optimized formulation for a liquid
after treatment with 0.5% boric acid for 3 wk. Linepithema humile bait, along with a unique, efficient bait station, to reduce the D.
demonstrated &90% worker mortality after exposure to 0.5% boric thoracicus worker population in a laboratory setting. Additional
acid for 2 wk (Hooper-Bui and Rust 2000). In addition, at higher studies are required to determine whether this bait can satisfactorily
concentration of toxicants, the bait solution may have a repellent ef- control D. thoracicus population in the field.
fect on target ants. Compared with that containing 10% sucrose,
the bait with 5% boric acid was repellent to S. invicta (Klotz et al.
1997). In L. humile, consumption significantly decreased after treat- References Cited
ment with baits containing 2% or 4% boric acid compared with Abbott, W. 1925. A method of computing the effectiveness of an insecticide. J.
that after treatment without boric acid (Klotz et al. 2000). Here, 3% Econ. Entomol. 18: 265267.
boric acid or borax bait solutions demonstrated no repellency Bluthgen, N., and K. Fiedler. 2004. Preferences for sugars and amino acids
against D. thoracicus, suggesting that D. thoracicus may have a and their conditionality in a diverse nectar-feeding ant community. J. Anim.
lower awareness of boric acid than L. humile does. Therefore, a bait Ecol. 73: 155166.
Bluthgen, N., G. Gottsberger, and K. Fiedler. 2004. Sugar and amino acid
effective in controlling one pest should not be applied for another
composition of ant-attended nectar and honeydew sources from an
pest without evaluation.
Australian rainforest. Austral Ecol. 29: 418429.
In the field, the effectiveness of baits in controlling pest ants may Buczkowski, G., E. Roper, and D. Chin. 2014. Polyacrylamide hydrogels: An
change with the season. In the wood ant species Formica lugubris effective tool for delivering liquid baits to pest ants (Hymenoptera:
Zetterstedt, the median effective concentration (ED50) of sucrose so- Formicidae). J. Econ. Entomol. 107: 748757.
lution is up to 0.9 M during late June, whereas it decreases to 0.4 M Cassill, D. L., and W. R. Tschinkel. 1999. Regulation of diet in the fire ant,
during early November (Sudd and Sudd 1985). Stein et al. (1990) Solenopsis invicta. J. Insect Behav. 12: 307328.
observed that S. invicta prefers carbohydrate baits in January and Cherrett, J. M., and T. Lewis. 1974. Control of insects by exploiting their be-
February. By contrast, more foragers select protein baits, rather than havior, pp. 130146. In D. F. Williams (ed.), Exotic ants Biology, impact
carbohydrate baits, from March to October in Houston, TX. and control of introduced species. Blackwell Scientific, Oxford, England.
Chong, K.-F., and C.-Y. Lee. 2009. Evaluation of liquid baits against field
Moreover, optimal bait density is critical for the effective and cost-
populations of the longlegged ant (Hymenoptera: Formicidae). J. Econ.
saving management of pests in the field (Nelson and Daane 2007).
Entomol. 102: 15861590.
Toxicants may become concentrated in bait solutions because of Cox, D. R., and D. Oakes. 1984. Analysis of survival data, vol. 21, Chapman
water evaporation, indirectly interfering with their acceptance by & Hall, London, United Kingdom.
ants (Klotz et al. 2004). Thus, our next step will be to examine Daane, K. M., L. Cooper, K. R. Sime, E. H. Nelson, M. C. Battany, and M. K.
whether our bait solution is effective in controlling D. thoracicus in Rust. 2008. Testing baits to control Argentine ants (Hymenoptera:
the field. Formicidae) in vineyards. J. Econ. Entomol. 101: 699709.
8 Journal of Economic Entomology, 2017, Vol. 0, No. 0

Dussutour, A., and S. Simpson. 2008. Carbohydrate regulation in relation to Nelson, E. H., and K. M. Daane. 2007. Improving liquid bait programs for
colony growth in ants. J. Exp. Biol. 211: 22242232. Argentine ant control: Bait station density. Environ. Entomol. 36:
Edwards, J., and L. Abraham. 1990. Changes in food selection by workers of 14751484.
the Pharaohs ant, Monomorium pharaonis. Med. Vet. Entomol. 4: Nishida, G. 2002. Hawaiian Terrestrial Arthropod Checklist, 4th ed. Bishop
205211. Museum Press, Honolulu, HI.
Fischer, M., and A. Shingleton. 2001. Host plant and ants influence the honey- Quarles, W. 1992. Borates provide least-toxic wood protection. IPM
dew sugar composition of aphids. Funct. Ecol. 15: 544550. Practitioner. Online.
Grover, C. D., A. D. Kay, J. A. Monson, T. C. Marsh, and D. A. Holway. Raymond, M. 1985. Presentation dune programme danalyse logprobit pour
2007. Linking nutrition and behavioural dominance: Carbohydrate scarcity microordinateur. Cah. ORSTOM Ser. Entomol. Med. Parasitol. 22:
limits aggression and activity in Argentine ants. Proc. R. Soc. Lond. B Biol. 117121.
Sci. 274: 29512957. Reimer, N. 1994. Distribution and impact of alien ants in vulnerable
Hahn, D. A., and D. E. Wheeler. 2002. Seasonal foraging activity and bait Hawaiian ecosystems, pp. 1122. In D. F. Williams (ed.), Exotic ants: biol-
ogy, impact, and control of introduced species. Westview Press, Boulder,
preferences of ants on Barro Colorado Island, Panama. Biotropica 34:
CO.
348356.
Ricks, B., and B. S. Vinson. 1970. Feeding acceptability of certain insects and
Hara, A. H., K. L. Aoki, S. K. Cabral, and R. Niino-DuPonte. 2014.
various water-soluble compounds to two varieties of the imported fire ant.
Attractiveness of gel, granular, paste, and solid formulations of ant bait in-
J. Econ. Entomol. 63: 145148.
secticides to the little fire ant, Wasmannia auropunctata (Roger)
Riley, C. 1889. The little red ant. Insect Life 2: 106108.
(Hymenoptera: Formicidae). Proc. Hawaii. Entomol. Soc. 46: 4554.
Rust, M. K., K. Haagsma, and D. A. Reierson. 1996. Barrier sprays to control
Ho, C., and K. Khoo. 1997. Partners in biological control of cocoa pests:
Argentine ants (Hymenoptera: Formicidae). J. Econ. Entomol. 89: 134137.
Mutualism between Dolichoderus thoracicus (Hymenoptera: Formicidae)
Rust, M. K., D. A. Reierson, and J. H. Klotz. 2004. Delayed toxicity as a crit-
and Cataenococcus hispidus (Hemiptera: Pseudococcidae). Bull. Entomol.
ical factor in the efficacy of aqueous baits for controlling Argentine ants
Res. 87: 461470. (Hymenoptera: Formicidae). J. Econ. Entomol. 97: 10171024.
Hooper-Bui, L. M., and M. K. Rust. 2000. Oral toxicity of abamectin, boric Silverman, J., and H. R. Tai. 2001. Acceptance and intake of gel and liquid
acid, fipronil, and hydramethylnon to laboratory colonies of Argentine ants sucrose compositions by the Argentine ant (Hymenoptera: Formicidae). J.
(Hymenoptera: Formicidae). J. Econ. Entomol. 93: 858864. Econ. Entomol. 94: 511515.
Kaplan, E. L., and P. Meier. 1958. Nonparametric estimation from incomplete Silverman, J., and R. J. Brightwell. 2008. The Argentine ant: Challenges in
observations. J. Am. Stat. Assoc. 53: 457481. managing an invasive unicolonial pest. Annu. Rev. Entomol. 53: 231252.
Kaspari, M., and S. P. Yanoviak. 2001. Bait use in tropical litter and canopy Smith, F. 1860. Descriptions of new species of hymenopterous insects collected
ants-evidence of differences in nutrient limitation. Biotropica 33: 207211. by Mr. AR Wallace at Celebes. Zool. J. Linn. Soc. 5: 5793.
Khoo, K., and G. Chung. 1989. Use of the black cocoa ant to control mirid Smith, L., A. Appel, and G. Keever. 1996. Cockroach control methods can
damage in cocoa. Planter 65: 370383. cause other pest problems. Ala. Agric. Exp. Stn. Highlights Agric. Res. 43:
Khoo, K., and C. Ho. 1992. The influence of Dolichoderus thoracicus 56.
(Hymenoptera: Formicidae) on losses due to Helopeltis theivora Sola, F., A. Falibene, and R. Josens. 2013. Asymmetrical behavioral response
(Heteroptera: Miridae), black pod disease, and mammalian pests in cocoa in towards two boron toxicants depends on the ant species (Hymenoptera:
Malaysia. Bull. Entomol. Res. 82: 485491. Formicidae). J. Econ. Entomol. 106: 929938.
Klotz, J. H., and J. I. Moss. 1996. Oral toxicity of boric acid-sucrose water Stein, M. B., H. G. Thorvilson, and J. W. Johnson. 1990. Seasonal changes in
bait to Florida carpenter ants (Hymenoptera: Formicidae). J. Entomol. Sci. bait preference by red imported fire ant, Solenopsis invicta (Hymenoptera:
31: 912. Formicidae). Fla. Entomol. 73: 117123.
Klotz, J. H., and D. Williams. 1996. New approach to boric baits. IPM Pract. Sudd, J., and M. Sudd. 1985. Seasonal changes in the response of wood-ants
18: 14. (Formica lugubris) to sucrose baits. Ecol. Entomol. 10: 8997.
Klotz, J. H., K. M. Vail, and D. F. Willams. 1997. Toxicity of a boric acid- Sumida, S., E. C. Da Silva-Zacarin, P. Decio, O. Malaspina, F. C. Bueno, and
sucrose water bait to Solenopsis invicta (Hymenoptera: Formicidae). J. O. C. Bueno. 2010. Toxicological and histopathological effects of boric
Econ. Entomol. 90: 488491. acid on Atta sexdens rubropilosa (Hymenoptera: Formicidae) workers. J.
Klotz, J. H., M. K. Rust, and P. Phillips. 2004. Liquid bait delivery systems for Econ. Entomol. 103: 676690.
controlling Argentine ants in citrus groves (Hymenoptera: Formicidae). Taylor, R. W. 1987. A checklist of the ants of Australia, New Caledonia and
New Zealand (Hymenoptera: Formicidae), vol. 41, C.S.I.R.O Division of
Sociobiology 43: 419427.
Entomology Report.
Klotz, J. H., D. H. Oi, K. M. Vail, and D. F. Williams. 1996. Laboratory evalu-
Ulloa-Chac! on, P., and G. I. Jaramillo. 2003. Effects of boric acid, fipronil,
ation of a boric acid liquid bait on colonies of Tapinoma melanocephalum
hydramethylnon, and diflubenzuron baits on colonies of ghost ants
Argentine ants and Pharaoh ants (Hymenoptera: Formicidae). J. Econ.
(Hymenoptera: Formicidae). J. Econ. Entomol. 96: 856862.
Entomol. 89: 673677.
Van Mele, P., and N. T. Cuc. 2001. Farmers perceptions and practices in use
Klotz, J. H., L. Greenberg, C. Amrhein, and M. K. Rust. 2000. Toxicity and
of Dolichoderus thoracicus (Smith) (Hymenoptera: Formicidae) for biolo-
repellency of borate-sucrose water baits to Argentine ants (Hymenoptera:
gical control of pests of sapodilla. Biol. Control 20: 2329.
Formicidae). J. Econ. Entomol. 93: 12561258.
Vander Meer, R., C. Lofgren, and J. Seawright. 1995. Specificity of the red im-
Knight, R. L., and M. K. Rust. 1990. Repellency and efficacy of insecticides
ported fire ant (Hymenoptera: Formicidae) phagostimulant response to
against foraging workers in laboratory colonies of Argentine ants carbohydrates. Fla. Entomol. 144154.
(Hymenoptera: Formicidae). J. Econ. Entomol. 83: 14021408. Wada, A., Y. Isobe, S. Yamaguchi, R. Yamaoka, and M. Ozaki. 2001. Taste-
Lanza, J., E. L. Vargo, S. Pulim, and Y. Z. Chang. 1993. Preferences of the fire enhancing effects of glycine on the sweetness of glucose a gustatory aspect
ants Solenopsis invicta and S. geminata (Hymenoptera: Formicidae) for of symbiosis between the ant, Camponotus japonicus, and the larvae of the
amino acid and sugar components of extrafloral nectars. Environ. Entomol. lycaenid butterfly, Niphanda fusca. Chem. Senses 26: 983992.
22: 411417. Ward, D. F., J. R. Beggs, M. N. Clout, R. J. Harris, and S. OConnor. 2006.
Lee, C.-Y., and L.-C. Lee. 2002. Field and laboratory evaluation of a boron- The diversity and origin of exotic ants arriving in New Zealand via human-
based containerized dual-bait formulation against the Pharaoh ant, mediated dispersal. Divers. Distrib. 12: 601609.
Monomorium pharaonis (Hymenoptera: Formicidae). Sociobiology 40: Way, M., and B. Bolton. 1997. Competition between ants for coconut palm
655665. nesting sites. J. Nat. Hist. 31: 439455.

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