Open access journal: http://periodicos.uefs.br/ojs/index.php/sociobiology
ISSN: 0361-6525

DOI: 10.13102/sociobiology.v60i4.453-458Sociobiology 60(4): 453-458 (2013)

Evaluation of formic acid toxicity to subterranean termite, Reticulitermes chinensis Snyder

YJ Xie, Q Du, QY Huang, CL Lei

Introduction

Formic acid is the simplest organic acid, present in 
many organisms. It is found in most plant species, including 
Polygonum hydropiper, Urttca dioica, Urtica urens (Hutchens 
1992, Chopra & Chopra 2006, Rahmatullah et al. 2010). In 
animal, formic acid is the major component of ants from the 
subfamily Formicinae. The produced formic acid concentra-
tion in such ant produced substances can reach up to 54% 
(Hölldobler & Wilson 1990). Formic acid also occurs in cara-
bid beetles (Will et al. 2010, Rossini et al. 1997, Attygalle et 
al. 1992) and notodontid caterpillars (Weatherston et al. 1979, 
Attygalle et al. 1993).

Formic acid was reported to possess significant insec-
ticidal activity. It has been used as a fumigant to control the 
varroa mite, Varroa destructor Anderson (Sharma et al. 1983, 
Underwood & Currie 2004, 2005, 2007, vanEngelsdorp et al. 
2008, Calderone 2010, Giovenazzo & Dubreuil 2011), and 
the tracheal mite, Acarapis woodi Rennie (Hoppe et al. 1989, 
Wilson et al. 1993, Nelson et al. 1994), and red imported fire 

Abstract
This study examined formic acid contact and fumigation toxicity to Reticulitermes 
chinensis in the laboratory. In the contact toxicity bioassay, the LD

50
 values ranged 

from 267.86 to 287.68 μg adult-1 for workers, 279.09 μg adult-1 for alates (male and 
female) and 223.08 μg adult-1 for soldiers after 24 h, respectively. In the fumigation 
bioassays, the LC

50
 values ranged from 0.84 to 1.08 μg ml-1 for workers, 1.19 μg ml-1 

for alates (male and female) and 0.57 μg ml-1 for soldiers after 24 h, respectively. At 
the concentration of 2.50 μg ml-1, the KT

50
 value of formic acid ranged from 25.38 to 

34.75 min for workers, from 42.21 to 45.62 min for alates (male and female), from 
32.18 to 36.37 min for soldiers. Although formic acid was significantly less toxic to 
subterranean termite than bifenthrin, but higher toxic to many other pest insects. 
The findings of this study provide important confirmation of formic acid with fumi-
gation toxicity against termite. It may be worth investigating the use of formic acid 
for managing subterranean termite.

Sociobiology
An international journal on social insects

Huazhong Agricultural University, Wuhan, China.

rESEArCH ArTICLE - TErMITES

Article History
Edited by 
Alexandre Vasconcellos, UFPB, Brazil
received       02 April 2013
Initial acceptance      19 May 2013
Final acceptance      18 June 2013

Keywords: Formic acid; contact toxicity; 
fumigation toxicity 

Corresponding author
Chao Liang Lei 
Hubei Insect resources Utilization and 
Sust. Pest Manag. Key Laboratory
Huazhong Agricultural University,
Wuhan 430070, China
E-Mail: cl_lei@yahoo.com.cn 
            ioir@mail.hzau.edu.cn

ants, Solenopsis invicta Buren (Chen et al. 2012), and stored 
product insects, Sitophilus oryzae and Rhyzopertha domini-
ca (Chaskopoulou 2007), and the mosquito larvicidal, Culex 
quinquefasciatus and Aedes aegypti (Welling & Paterson 
1985), and Drosophila and houseflies (Song & Scharf 2008a, 
b, 2009). 

Termites are world-wide pests threatening agriculture 
and the urban environment (Verma et al. 2009) worldwide. 
They cause over 3 billion dollars worth of damage to woo-
den structures annually throughout the U.S. (Su & Scheffrahn 
1998). Control and repair costs due to termites in China, have 
been increasing annually. Reticulitermes chinensis Snyder 
(Isoptera: Rhinotermitidae) is a species of termites with broad 
distribution that damages the wooden structures of buildings 
and the xylems of living old trees in China (Wei et al. 2007). 
Over the past two decades, the control of termite was usu-
ally accomplished by using organochlorines and organophos-
phates, which have been banned owing to environmental and 
human health concerns (Potter 1997). At the present time, se-
veral termiticides, including bifenthrin, chlorfenapyr, cyper-



YJ Xie, Q Du, QY Huang, CL Lei - Formic acid toxicity to Reticulitermes chinensis454

methrin, fipronil, imidacloprid and permethrin, were registe-
red for termite control around the world (UNEP Report 2000). 
However, the persistent use of synthetic chemicals to control 
termites raises several concerns related to environment and 
human health. In order to reduce the negative impact of pesti-
cides on the environment and to minimize the development of 
insecticide resistance in target pest species, as a result, the se-
arch for good efficacy and environmentally friendly insectici-
de is essential. As a safer or ‘green’ alternative, plant derived 
compounds have been used in termite management as contact 
insecticides or repellents, such as β-Thujaplicin, Thujopsene, 
T-muurolol, Cinnamaldehyde and Nootkatone (Nakashima 
& Shimizu 1972, Yoshida et al. 1998, Maistrello et al. 2001, 
Chang & Cheng 2002, Cheng et al. 2004). Another potential 
source of natural toxins against termite is the defensive/offen-
sive chemicals in ants. Some genus such as Leptogenys, Cen-
tromyrmex, Termitopone, Megaponera and Pheidole are all 
obligate predators of termites (Hölldobler & Wilson 1990). In 
addition, Kenne et al. (2000) observed that Myrmicaria opa-
civentris can quickly subdued Macrotermes bellicosus soldier 
with its pretarsus and abdominal thorn, thereby attacking 
the workers. Nascimento (2001) reported that the Pheidole 
pallidula preyed on Reticulitermes lucifugus, which play an 
important role in suppressing the establishment of termite co-
lonies. Pachycondyla analis, living in the dry forests of the 
Tanzanian coast and feeding on termites, can attack the termi-
te nest (Bayliss & Fielding 2002). Ke et al. (2008) found that 
Diacamma rugosum, Harpegnathos venator, Pachycondyla 
astuta and Polyrhachis dives had strong agonistic behavior 
and attacking capabilities to C. formosanus, and also obser-
ved the ant nest adjacent to the O. formosanus nest and the 
ant predation behavior to termite in the wild. The defensive 
chemicals of ants how to affect termite is not well understood, 
let alone the utilization of those chemicals in termite control. 
Therefore, the aim of present work was to study the contact 
and fumigation toxicity of formic acid to R. chinensis in the 
laboratory.

Methods

Termite

The termite samples of R. chinensis colonies were col-
lected from Huazhong Agriculture University, Wuhan, China. 
Colonies has been reared on wood pieces in a glass container 
(70 cm × 40 cm × 30 cm) in dark at 26 ± 1ºC and 80 ± 5% 
relative humidity for more than six months.

Contact toxicity

Contact toxicity of formic acid (98%, Sinopharm che-
mical Reagent Co., Ltd. China) to R. chinensis workers, ala-
tes (male and female) and soldiers was determined. For the 
ease of handling and obtaining uniform body weight, large 

workers were selected in the bioassay. Acetone was used as 
solvent, and the solution was applied with a 1.0 μl capilla-
ry tube. For workers, alates (male and female) and soldiers, 
doses of 100, 150, 200, 250, 300 and 350 μg adult-1 were ap-
plied. Controls were treated with acetone. Twenty adults of R. 
chinensis workers, alates (male and female) and soldiers were 
used for each concentration and control, and the experiment 
was replicated 3 times. Four colonies were used for workers. 
For alates (male and female) and soldiers, only one colony 
was tested. The contact toxicity of bifenthrin (98%, Huang-
ma Agrochemicals Co., Ltd, Jiangsu, China) to R. chinensis 
workers was also estimated using 2 colonies. Doses of 3.125, 
6.25, 12.5, 25.0 and 50.0 ng adult-1 were used. Bifenthrin is 
the most widely used commercial termiticide as positive con-
trol. Each dose was replicated 3 times, and each replicate con-
sisted of 20 workers. Both treated and control insects were 
then transferred to glass Petri dishes, and kept under the same 
environmental conditions described for the rearing. Mortality 
percentages were recorded after treatment for 24 h and LD50 
values were calculated according to Finney (1971).

Fumigation toxicity

To determine the fumigant toxicity of formic acid and 
the median effective time to cause mortality in 50 % of the 
workers from four different colonies (LT50 values), filter pa-
pers (Whatman No 1, cut into 2 cm diameter pieces) were 
impregnated with an appropriate concentration of 1.25, 2.5, 
5.0, 10.0 and 20.0 μg ml-1, respectively. The impregnated fil-
ter paper was then attached to the undersurface of the 1000 
ml glass jar’s (10 cm in diameter × 12.5 cm in height) screw 
cap, respectively. The cap was tightly screwed onto the jar, 
which contained 20 termites. In addition, workers from two 
other colonies were tested only at 2.5 μg ml-1. Alates (male 
and female) and soldiers were also tested at 2.5 μg ml-1. Two 
colonies were used for alates (male and female) and soldiers. 
Each concentration and control was replicated three times. 
Mortality was counted every minute after the formic acid 
had been delivered into the tube until all termites were dead. 
When no leg or antennal movements were observed, insects 
were considered dead.

Another experiment was designed in order to determi-
ne the 50% lethal concentration. For the mortality bioassay, 
20 workers, 20 alates (male and female) or 20 soldiers were 
placed in the bottle. The number of dead termites was coun-
ted after the formic acid had been delivered into the bottle 
for 24 h. Six concentrations were tested, including 0.20, 0.40, 
0.60, 0.80, 1.0 and 1.2 μg ml-1 for workers, alates (male and 
female) and soldiers, respectively. Three colonies were used 
for workers, and only one colony was used for alates (male 
and female) and soldiers. For each concentration × colony 
combination, there were three replicates. All bioassays were 
conducted at 26 ± 1ºC and 80 ± 5% relative humidity. The 
fumigation toxicity of bifenthrin (Huangma Agrochemicals 



Sociobiology 60(4): 453-458 (2013) 455

Co., Ltd, Jiangsu, China) on workers was also measured. Six 
concentrations were tested, including 0.125, 0.25, 0.625, 1.25 
and 2.5 ng ml-1. Two colonies were used, and there were three 
replicates for each colony.

Data analysis

Polo Plus v.1.0 (LeOra Software, Petaluma, CA) was 
used to estimate LD50, KT50 and LC50 with 95% confidence 
intervals (CIs). The relative toxicity ratio with their upper and 
lower 95% confidence limits was used to evaluate the signifi-
cance of the difference between LD50, KT50 and LC50 values. 
The significance was set at the P = 0.05 probability level. If 
the 95% confidence interval of the ratio between two LD50, 
KT50 or LC50 values included 1, these were not considered to 
be significantly different (Robertson et al. 2007).

Results

Toxicity of formic acid against R. chinensis was shown 
in Table 1. The LD50 values of formic acid were ranging from 
267.86 to 287.68 μg adult-1 for workers, which were larger 
than those of bifenthrin with LD50 values ranging from 7.16 to 
10.39 ng adult-1, and no significant difference was also found 
in LD50 values of formic acid for workers between colony A 
and the other three colonies. Alates and soldiers were more 
sensitive to formic acid than workers. The LD50 values for ala-
tes (male and female) and soldiers were at 279.09 and 223.08 
μg adult-1, respectively.

Discussion

When applied topically, formic acid was significantly 
less toxic to subterranean termite than bifenthrin, but higher 
toxic to many other pest insects. The LC50 value of formic 
acid for rice weevil, Sitophilus oryzae, was 6.03-7.60 μg ml-1, 
and for the mosquito, Culex quinquefasciatus, was 3.67 μg 
ml-1 (Welling & Paterson 1985). Formic acid displayed low 
toxicity against yellow fever mosquito, Aedes aegypti, at 6.00 
μg ml-1 (Chaskopoulou 2007). As demonstrated recently by 
Chen et al. (2012), the LC50 value of formic acid was 0.26-
0.70 μg ml-1 for red imported fire ants. This indicated that the 
toxicity of formic acid to subterranean termite, R. chinensis, 
was higher than that of the insects studied so far, but similar 
to red imported fire ants.

a W: works; A: alates (male and female); S: soldiers.
b Positive control.

As can be seen from Table 2, the LC50 values ranged 
from 0.84 to 1.08 μg ml-1 for workers, 1.19 μg ml-1 for alates 
(male and female) and 0.57 μg ml-1 for soldiers after treat-
ment for 24 h, respectively. The LC50 values of a well-known 
commercial pesticide used as a positive control in this study, 
bifenthrin, were 0.40 and 0.42 ng ml-1, respectively (Table 2). 
At a concentration of 2.50 μg ml-1, the KT50 value of formic 
acid ranged from 25.38 to 34.75 min for workers, from 42.21 
to 45.62 min for alates (male and female), from 32.18 to 36.37 
min for soldiers (Table 3). Meanwhile, the higher the concen-
tration, the smaller the KT50 value was. Within A colony, the 
KT50 value of workers was always significantly smaller than 
that of alates (male and female), and soldiers.

Table 1- LD50 values of formic acid by contact against workers, ala-
tes and soldiers of R. chinensis.

Table 2- LC50 values of formic acid applied as fumigant for 24 h 
against workers, alates and soldiers of R. chinensis.

a W: works; A: alates (male and female); S: soldiers.
b Positive control.

Table 3- KT50 values of formic acid applied as fumigant against 
workers of R. chinensis.

The action mode of formic acid was to inhibit the ac-
tivity of mitochondrial cytochrome c oxidase (Nicholls 1975, 
Petersen 1977). Recently, formic acid has been reported to 
have a significant excitatory effect on the nervous system of 
housefly larvae (Song & Scharf 2008a, b). Song & Scharf 
(2009) reported the formic acid have a significant excitatory 
effect on the nervous system of Drosophila melanogaster. It 
may explain why formic acid has such a low contact toxicity 

a W: works; A: alates (male and female); S: soldiers.

Chemical Colony Caste a Slope (±SE) LD50 95% CI
Formic
Acid

Bifenthrin b

A
B

C
D
E
E
F

W
W
A
W
W
S
W
W

2.36(±0.31) 
2.09(±0.29)
2.91(±0.35)
2.54(±0.32)
2.21(±0.29)
2.93(±0.31)
2.52(±0.26)
2.46(±0.24)

287.68 (μg adult-1)
286.24 (μg adult-1)
279.09 (μg adult-1)
273.77 (μg adult-1)
267.86 (μg adult-1)
223.08 (μg adult-1)
7.16 (ng adult-1)
10.39 (ng adult-1)

250.02-347.80
245.04-322.94
248.91-250.48
241.01-322.79
232.15-322.98
200.83-250.48

5.99-8.41
8.82-12.20

Chemical Colony Caste a Slope (±SE) LC50 95% CI
Formic
Acid

Bifenthrin b

A
C
B

C
D

W
W
W
A
S
W
W

6.71(±0.76)
4.56(±0.52)
4.39(±0.59)
4.72(±0.73)
4.77(±0.54)
3.06(±0.29)
2.80(±0.26)

0.95 μg ml-1
0.84 μg ml-1
1.08 μg ml-1
1.19 μg ml-1
0.57 μg ml-1
0.40 ng ml-1
0.42 ng ml-1

0.88-1.03
0.77-0.93
0.98-1.23
1.07-1.38
0.49-0.65
0.32-0.50
0.35-0.50

Colony ID Caste a Dosage (μg/ml) Slope (±SE) KT50 (min) 95% CI
A

B

C

D

E

F

W
W
W
W
W
W
W
W
W
W
A
S
W
W
W
W
W
W
W
W
W
W
W
A
S
W

1.25
2.5
5.0

10.0
20.0
1.25
2.5
5.0

10.0
20.0
2.5
2.5

1.25
2.5
5.0

10.0
20.0
1.25
2.5
5.0

10.0
20.0
2.5
2.5
2.5
2.5

4.32(±0.77)
7.87(±0.75)
9.06(±1.00)
9.63(±1.43)
5.10(±0.58)
2.32(±0.28)
8.45(±0.81)
9.47(±0.88)
6.24(±0.64)
7.15(±0.72)
7.77(±1.06)
4.19(±0.51)
9.84(±0.84)
7.95(±0.63)
11.32(±0.96)
14.18(±1.37)
15.62(±1.63)
14.05(±1.35)
12.03(±1.13)
6.02(±0.66)
8.56(±0.79)
5.72(±0.64)
12.94(±1.24)
9.40(±1.47)
8.30(±1.44)
6.95(±0.62)

91.67
26.71
16.79
8.74
5.14

107.45
32.85
19.70
9.55
4.74

45.62
36.37
84.12
25.38
19.63
7.74
5.25

79.53
29.74
17.90
6.31
3.79

34.75
42.21
32.18
28.25

82.27-109.12
25.47-28.01
15.80-17.82

7.78-9.34
3.87-5.48

94.26-123.01
31.61-34.25
18.97-20.49
9.05-10.05
4.42-5.03

43.34-48.79
33.21-41.01
81.25-86.94
24.21-26.51
19.05-20.20

7.44-8.04
5.08-5.42

76.53-82.67
28.49-31.07
16.54-19.72

5.94-6.68
3.01-4.54

33.52-36.02
40.14-45.72
30.30-35.30
26.44-29.90



YJ Xie, Q Du, QY Huang, CL Lei - Formic acid toxicity to Reticulitermes chinensis456

compared with fumigation.
Fumigation has been extensively applied as building 

treatment in termite management programs. Fumigants in-
clude methyl bromide, sulfuryl fluoride, dichloroethane and 
dibromoethane. However, compared to contact insecticides, 
fumigation has lower toxicity. Therefore, the high efficacy 
of the conventional synthetic contact insecticides may have 
slowed down the development of any alternatives.

Using formic acid directly as fumigant against subter-
ranean termite may not be suitable for its own acidity and 
corrosiveness. However, formate ester is much less corrosive 
than formic acid and should be much easier to handle. Scharf 
et al. (2006) and Nguyen et al. (2007) also found the low mo-
lecular weight formate esters are volatile compounds with 
fumigant insecticidal activity. Similarly, Chaskopoulou et al. 
(2009) reported the volatile low molecular weight compounds 
against Aedes aegypti and Culex quinquefasciatus. Previous 
research reported some formate esters had neurological activ-
ity on Drosophila melanogaster Meig and Musca domestica 
L, owning to their hydrolyzed metabolite, formic acid (Song 
& Scharf 2008a, b). Previously, Haritos & Dojchinov (2003) 
reported some formate esters exert toxicity in the stored prod-
uct beetle Sitophilus oryzae L. for its hydrolyzed metabolite, 
formic acid, which causes mitochondrial impacts. Therefore, 
formate esters may also be as good candidates for subterra-
nean termite, particularly as fumigant in treatment. However, 
the prerequisite is that formic acid can be liberated from for-
mate esters in termite bodies. This is definitely worth for fur-
ther investigation.

The formic acid fumigation bioassay to termite was 
carried out in an enclosed environment for termite control, 
such as house, storage spaces and timber. In order to make fu-
migation effectively, the enclosed environment must be intact 
during the application. Simultaneously, other factors such as 
temperature, humidity, the concentration and time of formic 
acid applied all should be considered. These are the key of 
successful application of formic acid in the termite control. 
In addition to considering their potential fumigant effect, like 
other commercial fumigant, the more important concern of 
formic acid application is its toxicity to humans and the envi-
ronment. The current Occupational Safety and Health Admin-
istration (OSHA) permissible exposure limit (PEL) is 3 mg 
m-3 as the 8 h time-weighted average (TWA) concentration 
for DDVP, 9 mg m-3 for formic acid, 80 mg m-3 for methyl 
bromide, 250 mg m-3 for methyl formate and ethyl formate 
is 300 mg m-3 (Chen et al. 2012). Therefore, the effect of for-
mate esters against termite is more worth to investigate, as it 
seems that formate esters may be more practical to use than 
formic acid.

Acknowledgments

This study was supported by the Fundamental Re-
search Funds for the Central Universities (2010PY065).

References

Attygalle A. B., J. Meinwald & T. Eisner 1992. Defensive se-
cretion of a carabid beetle, Helluomorphoides clairvillei. J. 
Chem. Ecol., 18: 489-498.

Attygalle, A. B., S. R. Smedley, J. Meinwald & T. Eisner 1993. 
Defensive secretion of two notodontid caterpillars (Schizura 
unicornis, S. badia). J. Chem. Ecol.,  19: 2089-2104.

Bayliss, J. & A. Fielding 2002. Termitophagous foraging by 
Pachycondyla analis (Formicidae: Ponerinae) in a Tanzanian 
coastal dry forest. Sociobiology, 39: 103-122.

Calderone, N. W. 2010. Evaluation of Mite-Away-II for fall 
control of Varroa destructor (Acari: Varroidae) in colonies of 
the honey bee Apis mellifera (Hymenoptera: Apidae) in the 
northeastern USA. Exp. Appl. Acarol. 50: 123-132.

Chaskopoulou, A. 2007. Testing vapor toxicity of formate, 
acetate, and heterocyclic compounds to Aedes aegypti and 
Musca domestica. MS Thesis, University of Florida, Gaines-
ville, FL.

Chaskopoulou, A. S. Nguyen, R. M. Pereira, M. E. Scharf 
& P. G. Koehler 2009. Efficacy of 31 volatile low molecular 
weight compounds against Aedes aegypti and Culex quinque-
fasciatus. J. Med. Entomol., 46: 328-334.

Chang, S. T. & S. S. Cheng 2002. Antitermitic activity of 
leaf essential oils and components from Cinnamomum os-
mophleum. J. Agr. Food Chem., 50: 1389-1392.

Chen, J., T. Rashid & G. L. Feng 2012. Toxicity of formic 
acid to red imported fire ants, Solenopsis invicta Buren. Pest 
Manag. Sci. 68: 1393-1399.

Cheng, S. S., C. L. Wu, H. T. Chang, Y. T. Kao & S. T. Chang 
2004. Antitermitic and antifungal activities of essential oil 
of Calocedrus formosana leaf and its composition. J. Chem. 
Ecol.,  30: 1957-1967.

Chopra, R. N. & I. C. Chopra 2006. Indigenous Drugs of In-
dia. Bimal Kumar Dhur of Academic publishers, India.

Nascimento, F. S. 2001. Behavioral responses of Mediterra-
nean termite Reticulitermes lucifugus (Rossi) (Isoptera: Rhi-
notermitidae) under presence effects of ant Pheidole pallidula 
(Nyl.) (Hymenoptera: Formicidae). Rev. Bras. Zooc., 3: 195-
201.

Finney, D. J. 1971. Probit Analysis, 3rd edition. Cambridge 
University Press, Cambridge, UK.

Giovenazzo, P. & P. Dubreuil 2011. Evaluation of spring organic 
treatments against Varroa destructor (Acari: Varroidae) in honey 
bee Apis mellifera (Hymenoptera: Apidae) colonies in eastern 
Canada. Exp. Appl. Acarol., 55: 65-76.

Haritos, V. S. & G. Dojchinov 2003. Cytochrome c oxidase inhibi-
tion in the rice weevil Sitophilus oryzae by formate, the toxic metab-
olite of alkyl formates. Comp. Biochem. Physiol. C ,136: 135-143.



Sociobiology 60(4): 453-458 (2013) 457

Hölldobler, B. & E. O. Wilson 1990 The Ants. Cambridge, 
Mass. Harvard University Press, 559-569.

Hoppe, H., W. Ritter & E. W. C. Stephen 1989. The control of 
parasitic bee mites: Varroa jacobsoni, Acarapis woodi, and Tro-
pilaelaps clareae with formic acid. Am. Bee J., 129: 739-742.

Hutchens, A. R. 1992. A Handbook of Native American Herbs. 
Shambhala Publications, Massachusetts.

Ke, Y. L. T. Y. Zhuang, C. X. Wang, S. Zhao & W. J. Tian 
2008. Agonistic behavior of six ant species to Coptotermes 
formosanus (Isoptera: Rhinotermitidae) in laboratory assays. 
Sociobiology, 51: 199-206.

Kenne, M., A. Dejean, R. Fénéron & J. L. Durand 2000. 
Changes in worker polymorphism in Myrmicaria opaciventris 
Emery (Formicidae: Myrmicinae). Insect. Soc., 47: 50-55.

Maistrello, L., G. Henderson & R. A. Laine 2001. Effects of 
nootkatone and a borate compound on formosan subterranean 
termite (Isoptera: Rhinotermitidae) and its symbiont proto-
zoa. J. Entomol. Sci., 36: 229-236.

Meepagala, K. M., W. Osbrink, C. Burandt, A. Laxb & S. O. 
Dukea 2011. Natural-product-based chromenes as a novel 
class of potential termiticides. Pest Manag. Sci., 67: 1446-
1450.

Nakashima, Y., & K. Shimizu 1972. Antitermitic activity 
of Thujopsis dolabrata var Hondai. III. Components with a 
termiticidal activity. Miyazaki Daigaku Nogakubu Kenkyu 
Hokoku, 19: 251-259.

Nelson, D., P. Mills, P. Sporns, S. Ooraikul & D. Mole 1994. 
Formic acid application methods for the control of honey bee 
tracheal mites. Bee Sci., 3: 128-134.

Nicholls, P. 1975. Formate as an inhibitor of cytochrome c 
oxidase. Bioch. Biophys. Res. Commun., 67: 610-616.

Nguyen, S. N., C. Song & M. E. Scharf 2007. Toxicity, syn-
ergism, and neurological effects of novel volatile insecticides 
in insecticide susceptible and resistant Drosophila strains. J. 
Econ. Entomol., 100: 534-544.

Potter, M. F. 1997. Termites, in Handbook of Pest Control, 
8th edition, ed. by Hedges SA. Franzak & Foster, Cleveland, 
OH, pp. 233-333.

Petersen, L. C. 1977. The effect of inhibitors on the oxygen 
kinetics of cytochrome c oxidase. Bioch. Biophys. Acta, 460: 
299-307.

Rahmatullah, M., R. Jahan, M. S. Hossan, S. Seraj, M. M. 
Rahman, A. R. Chowdhury, Z. U. M. E. U. Miajee, D. Nasrin, 
Z. Khatun, F. I. Jahan & M. A. Khatun 2010. A Comparative 
Analysis of Medicinal Plants used by Several Tribes of Chit-
tagong Hill Tracts Region, Bangladesh to Treat Helminthic 
Infections. Adv. Nat. Appl. Sci., 4: 105-111.

Robertson, J. L., R. M. Russell, H. K. Preisler & N. E. Savin 

2007. Bioassays with Arthropods, 2nd edition. CRC Press, 
Boca Raton, FL.

Rossini, C., A. B. Attygalle, A. Gonzalez, S. R. Smedley, M. 
Eisner, J. Meinwald & T. Eisner 1997. Defensive production 
of formic acid (80 percent) by a carabid beetle (Galerita le-
contei). Proc. Nat. Acad. Sci. USA, 94: 6792-6797.

Scharf, M. E., S. N. Nguyen & C. Song 2006. Evaluation of 
volatile low molecular weight insecticides using Drosophila 
melanogaster as a model. Pest Manag. Sci., 62: 655-633.

Sharma, O. P., R. Garg & G. S. Dogra 1983. Efficacy of for-
mic acid against Acarapis woodi (Rennie). Indian Bee J., 45: 
1-2.

Song, C. & M. E. Scharf 2008a. Formic acid: a neurologi-
cally-active, hydrolyzed metabolite of insecticidal formate 
esters. Pest. Biochem. Physiol., 92: 77-82.

Song, C. & M. E. Scharf 2008b. Neurological disruption by 
low molecular weight compounds from the heterobicyclic and 
formate ester classes. Pest. Biochem. Physiol., 92: 92-100.

Song, C. & M. E. Scharf 2009. Mitochondrial impacts of in-
secticidal formate esters in insecticide-resistant and insecti-
cide-susceptible Drosophila melanogaster. Pest Manag. Sci., 
65: 697-703.

Su, N.Y. & R. H. Scheffrahn 1998. A review of subterranean 
termite control practices and prospects for integrated pest 
management programmes. Integr. Pest Manag. Rev., 3: 1-13.

Underwood, R. M. & R. W. Currie 2004. Indoor winter fu-
migation of Apis mellifera (Hymenoptera: Apidae) colonies 
with Varroa destructor (Acari: Varroidae) with formic acid is 
a potential control alternative in northern climates. J. Econ. 
Entomol., 97: 177-186.
Underwood, R. M. & R. W. Currie 2005. Effect of concentra-
tion and exposure time on treatment efficacy against Varroa 
mites (Acari: Varroidae) during indoor winter fumigation of 
honey bees (Hymenoptera: Apidae) with formic acid. J. Econ. 
Entomol., 98: 1802-1809.

Underwood, R. M. & R. W. Currie 2007. Effects of release 
pattern and room ventilation on survival of varroa mites and 
queens during indoor winter fumigation of honey bee colo-
nies with formic acid. Can. Entomol., 139: 881-893.

UNEP/FAO/Global IPM Facility Expert Group on Termite 
Biology and Management, 2000. Finding alternatives to per-
sistent organic pollutants (POPs) for termite management, on-
line at www.chem.unep.ch/pops/termites/termite_ch4.htm.
vanEngelsdorp, D., R. M. Underwood & D. L. Cox-Foster 
2008. Short-term fumigation of honey bee (Hymenoptera: 
Apidae) colonies with formic and acetic acids for the control 
of Varroa destructor (Acari: Varroidae). J. Econ. Entomol., 
101: 256-264.
Verma, M., S. Sharma & R. Prasad 2009. Biological alterna-



YJ Xie, Q Du, QY Huang, CL Lei - Formic acid toxicity to Reticulitermes chinensis458

tives for termite control: A review. Int. Biodeter. Biodegr., 63: 
959-972.

Weatherston J., J. E. Percy, L. M. MacDonald & J. A. Mac-
Donald 1979 Morphology of the prothoracic defensive gland 
of Schizura concinna (J.E. Smith) (Lepidoptera: Notodontidae) 
and the nature of its secretion. J. Chem. Ecol.,  5:165-177

Wei, J. Q., J. C. Mo, X. J. Wang & W. G. Mao 2007. Biol-
ogy and ecology of Reticulitermes chinensis Snyder (Isoptera: 
Rhinotermitidae) in China. Sociobiology, 50: 553-559.

Will K. W., A. S. Gill, H. Lee & A. B. Attygalle 2010. Quan-
tification and evidence for mechanically metered release of 
pygidial secretions in formic acid producing carabid beetles. 
J. Insect Sci., 10: 1-17.

Welling, W. & G. D. Paterson 1985. Toxicodynamics of in-
secticides, in Comprehensive Insect Physiology, Biochemis-
try and Pharmacology, Vol. 12, ed. by Kerkut GA and Gilbert 
LI. Pergamon, Oxford, UK, pp. 603-646.

Wilson, W.T., J. R. Baxter, A. M. Collins, R. L. Cox & T. D. 
Cardoso 1993. Formic acid fumigation for control of tracheal 
mites in honey bee colonies. Bee Sci., 3: 26-32.

Yoshida, S., Y. Morita, K. Narita & T. Okabe 1998. Termiti-
cidal efficacy of neutral oil obtained from Aomori hiba. Ten-
nen Yuki Kagobutsu Toronkai Koen Yoshishu, 40: 311-315.