J Pest Sci

DOI 10.1007/s10340-016-0778-z

ORIGINAL PAPER

Bait station preferences in two Macrotermes species

Naeem Iqbal1,2,3 • Lahiru S. Wijedasa1,4,5,6 • Theodore A. Evans1,7

Received: 7 December 2015 / Revised: 3 April 2016 / Accepted: 14 May 2016

 Springer-Verlag Berlin Heidelberg 2016

Abstract Baiting is considered to be a relatively environ-
mentally benign termite control method; however, all com-
mercial baiting systems are designed for species in the
Rhinotermitidae and are used primarily in temperate countries.
Fungus-growing termites in the Macrotermitidae can be
important pests in tropical countries; they can be difficult to
control using all available methods, and there are no baiting
systems designed for them. We tested bait station size, an
important component of bait station design, against two
Macrotermes species in Singapore. Macrotermes gilvus recrui-
ted to small stations (0.35 L) very poorly and medium stations
(3.6 L) poorly (both similar in size to various commercial sta-
tions), but they recruited to large stations (11.5 L) well.
Macrotermes carbonarius followed a similar pattern but
recruited to fewer stations overall. In the occupied stations, M.
gilvus ate the bait wood, sometimes creating a fungus garden
inside the stations, and brought little soil into the stations. In
comparison, M. carbonarius ate no wood at all, but filled stations
with soil. There was significantly less leaf litter around M. car-
bonarius mounds compared with M. gilvus mounds, although
there were no obvious differences in habitat, which suggested
that M. carbonarius eats leaf litter only and is not a pest species.
Our study shows that stations much larger than current com-
mercial options may provide a useful means for controlling pest
wood-eating, fungus-growing termites in tropical countries.
Keywords Bait size
Disturbance
Fungus-growing

Isoptera
Macrotermitinae
Termite

Communicated by M. Traugott.
Key messages
& Theodore A. Evans
cryptotermes@gmail.com • Baiting is a major control method for termites in
1
buildings in temperate countries, yet is not used against
National University of Singapore, Singapore 117345,
Singapore
fungus-growing pest termites in tropical countries.
2
• We tested bait station size and found large stations were
Department of Entomology, Faculty of Agricultural Sciences
and Technology, Bahauddin Zakariya University,
10 times more successful at attracting and retaining
Multan 60800, Pakistan termites than small stations, which were of similar size
3
Present Address: Department of Plant Protection, Faculty of
to commercial stations used in temperate countries.
Agricultural Sciences, Ghazi University, • Large stations could be used to develop baiting systems
Dera Ghazi Khan 32200, Pakistan against fungus-growing termites in the tropics, poten-
4
Singapore Botanic Gardens, 1 Cluny Road, tially including forestry and agriculture.
Singapore 259569, Republic of Singapore
5
Conservation Links, 433 Clementi Avenue 3, #01-258,
Singapore 120433, Singapore Introduction
6
Rimba, 18E Kampung Basung, 21700 Kuala Berang,
Terengganu, Malaysia Termites are major pests of wood and wood products,
7
Present Address: School of Animal Biology, University of
especially in tropical and subtropical areas of the world (Su
Western Australia, Perth, WA 6009, Australia and Scheffrahn 2000; Rouland-Lefèvre 2011). There are

123

various methods to prevent and control termite attack,
including physical barriers, chemical insecticide barriers in
soil and in building materials, dusting with insecticide
toxicants, and baiting (Edwards and Mill 1989; Pearce
1997; Hu 2011). Each method has various advantages
depending on the building construction type and density of
buildings, from free-standing wooden houses in low den-
sity suburbs to high rise concrete apartment blocks in the
high density inner city. Each method has differing aims,
from how they preventing the termites from causing
damage, to minimizing undesirable side effects, such as
negative environmental impacts.
Baiting was developed to reduce the population size of,
or even eliminate termite colonies, i.e. kill all the termites,
and to be more environmentally benign. The colony was
eliminated by feeding insecticides in a cellulose food matrix
to the termites, and this approach produced the environment
benefits: using less insecticide and containing the insecti-
cide inside a bait station (e.g. Esenther and Beal 1979; Su
1995; Grace et al. 1996; Evans 2006). Commercial baiting
systems were introduced around 20 years ago, and are now
the second most used termite control methods, at least in
wealthier countries (Evans and Iqbal 2015).
Current commercial baiting systems may not be useful
everywhere. Current baiting systems were designed in
temperate latitudes, especially in the USA, for use in urban
habitats, and to target pest termites in the Rhinotermitidae,
specifically species in Reticulitermes and Coptotermes (Su
1995; Grace et al. 1996). There are many other pest ter-
mites found in the tropics, in rural and forested habitats,
and in other termite families (Roonwal 1979; Cowie et al.
1989; Mitchell 2002; Constantino 2002). It is unclear
whether the current commercial baiting systems are
appropriate for these much wider range of circumstances;
in tropical Asia they are generally used only in urban
habitats against Coptotermes species (Lenz 2002; Lenz and
Evans 2002; Lee et al. 2007; Evans and Iqbal 2015).
One termite subfamily, the Macrotermitinae, may be of
particular concern: various species of Macrotermes, Mi-
crotermes, Odontotermes and other genera are important
pests in tropical Asia and Africa (Roonwal 1970; Wood
et al. 1980, 1987; Cowie et al. 1989; Tiben et al. 1990).
These termites do not have protozoan symbionts in their
guts, instead they cultivate a basidiomycete or ‘white rot’
fungus, Termitomyces spp, as a symbiont, thus their com-
mon name of ‘fungus-growers’ (Wood and Thomas 1989).
In general, they have larger body sizes, live in more pop-
ulous colonies, forage over longer distances, dominate
other termites living in their habitats, and are significant
components of these habitats (Evans et al. 1998; Bignell
and Eggleton 2000; Davies et al. 2003; Schuurman 2006;
Pringle et al. 2010). Fungus-growing termites may have
different requirements to the Rhinotermitidae; the
123
J Pest Sci
components of current commercial baits are still being
tested against them (e.g. Lee et al. 2014).
It seems likely that the major biological and ecological
differences between fungus-growing termites and the
rhinotermitid termites could drive different bait station
design. Food size and bait station design can have large
difference in rhinotermitid termite foraging behaviour
(Evans and Gleeson 2006; Lenz et al. 2009; Evans 2010).
In this study, we investigated the effect of bait station
design on two Macrotermes species in Singapore. We
considered the effect of size in attracting and retaining
foraging termites, as this is a critical attribute in the success
of any bait system (Evans and Iqbal 2015), and will need to
be optimized to make baiting systems work against fungus-
growing termites in tropical systems. We hypothesized that
larger bait stations with greater food resources would be
found more quickly, recruit more termites, and retain those
termites when disturbed during inspections.
Materials and methods
Study site
We performed the study at the Singapore Botanic Gardens
(SBG; Latitude 1.3120511N, Longitude 103.817417E).
The SBG is 74 hectares and contains a mixture of habitats,
including primary dipterocarp rainforest, various stages of
secondary forests, manicured parklands, and specialist
gardens. Mound-nests of two Macrotermes species,
Macrotermes gilvus and Macrotermes carbonarius, are
found in the forests and parkland, often near large trees.
We selected a total of 25 mound-nests: 17 M. gilvus and
eight M. carbonarius. The mounds were all considered to
contain healthy colonies, with complete clay covering,
recent construction activity, and termites foraging nearby.
The M. gilvus mounds were significantly smaller than those
of M. carbonarius mounds (t tests; height t = 1.989,
df = 21, p = 0.059; width t = 3.409, df = 21, p = 0.003;
Fig. 1). All these mounds were present near trees, and we
observed Macrotermes galleries in several trees, some of
which were culturally important and were under heritage
protection laws.
Bait stations
We tested three bait/monitoring station sizes. All were
constructed from cylindrical plastic containers with sloping
sides, with holes of 8 mm diameter, on sides and base, and
with a piece of wood in proportion to the size of the con-
tainer. The small stations were ca. 0.35 L (16.5 cm high,
top [ 8 cm, bottom [ 2.5 cm), and contained one piece of
J Pest Sci
250
Mound Diameter (cm)
200
150
100
50
0
0 20 40 60 80 100
Mound height (cm)
Fig. 1 Sizes of Macrotermes gilvus and M carbonarius mounds from
the Singaporean Botanic Gardens used in the study. Black
squares = M. gilvus; white circles M. carbonarius. Regressions M.
gilvus y = 0.8207x ? 47.098, r2 = 0.16; M. carbonarius y =
1.6084x ? 62.6, r = 2, R2 = 0.79
Pinus radiata wood (12.1 9 5.0 9 2.5 cm). This size was
comparable to the earliest commercial systems, including
Sentricon by Dow Chemical Company and Firstline by
FMC Corporation. The medium stations were ca 3.6 L in
volume (15.5 cm high, top [ 18.5 cm, bottom [ 16 cm),
and contained two pieces of wood (13.0 9 7.6 9 2.5 cm).
This size was comparable to later design bait systems,
including Exterra by Ensystex, Nemesis by PCT
International, and X-term by Sumitomo. The large sta-
tions were ca. 11.5 L (26.5 cm high, top [ 26.5 cm, bot-
tom [ 20.5 cm), contained four pieces of wood
(26.0 9 7.6 9 2.5 cm) placed vertically, with a fifth piece
(16.0 9 7.6 9 2.5 cm) placed on the base of the container.
Another piece of wood (26.5 9 15.2 9 2.5 cm) was
placed underneath each large station. The large station was
comparable in size to non-commercial monitoring stations
used in previous studies (Evans et al. 1998, 1999; Evans
and Gleeson 2001; Evans 2004, 2006; Lenz et al. 2012,
2013).
We installed four large, four medium, and four small
stations per mound for a total of 300 stations around the 25
mounds. We dug holes suitable in size to accommodate the
different-sized stations at ca. 2 m from the edge of the
mounds. We positioned the holes around the circle, so that
adjacent holes were 30 to each other, relative to the centre
of the mound, each hole and station ca. 1.5 m apart. We
place stations sequentially in size sets around the circle, i.e.
small, medium, large, small, medium, etc. The stations
were placed in the holes, covered with plastic lids (small [
12 cm, medium [ 21 cm, large [ 28 cm), on top of which
we placed concrete pavers and soil to prevent disturbance
from members of the public.
Data recording and analysis
We inspected all stations after 1, 2, 4, and 8 weeks. At each
inspection, we cleared the covering soil, moved the paver,
opened the lid, and visually checked for the presence of
termites or their mudding in each station. We attempted to
minimize disturbance of the stations. We would move
wood or mud as little as possible until we observed ter-
mites, and could identify the species. We would then
immediately replace the lid and paver, and cover with soil.
After the first week inspection, we recorded new contact,
continuous contact, and abandonment by the respective
Macrotermes species and other, non-target, termite species.
We defined ‘new’ contact to a station as the first appear-
ance of that species in that station, from inspection 2
onwards, and ‘continuous’ contact as the same species in
that station as observed in the previous inspection. We
defined ‘abandonment’ as the loss of a species in a station
in which it has been observed in the previous inspection.
We analysed data using repeated measures ANOVA,
because the units were sampled (i.e. inspected) repeatedly
(Sokal and Rohlf 1995). The ‘repeated’ factor was time,
with time interval adjusted for weeks (1, 2, 4 and 8) rather
than inspection number (1, 2, 3, and 4). As each inspection
of each station was not considered independent in the
analysis, there was no pseudoreplication. We performed all
analyses in SYSTAT (v. 9.0, 1998).
Results
Macrotermes gilvus
Total contacts to stations
The mean number of contacts by M. gilvus to stations
varied between station size and time. The mean number of
contacts to small stations was 0.53 stations/mound after
1 week, but this eventually decreased to 0.18 after eight
weeks. The mean number of contacts to medium stations
was similar to that of small stations after one week, 0.59
stations/mound, which remained steady until eight weeks,
0.65 stations/mounds. The mean number of contacts to
large size stations was similar to the small and medium
stations after 1 week, 0.47 stations/mound; however, this
increased fourfold to 1.94 stations/mound after 8 weeks
(Fig. 2). These combined differences were significant:
fewer small stations were occupied over time, whereas the
opposite was true for large stations (interaction
F6,144 = 10.155, p \ 0.001; station size F2,48 = 7.143,
p = 0.002; time F3,144 = 4.949, p = 0.003).
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 J Pest Sci

New contacts over time A

1.5

The mean number of new contacts to stations by M. gilvus

mimicked the pattern observed for total number of con- 1.0
tacts, but less extreme. The number of new contacts to
small stations fluctuated around the mean of 0.16 sta- 0.5
tions/mound over the 8 weeks of the experiment. The
number was similar, but slightly higher for medium sta- 0
tions: 0.22 stations/mound. However, the number of large B
stations contacted by M. gilvus increased from 0.53 to 0.94 1.5

Number of staons
over the experimental period (Fig. 3a). These combined
differences were significant: new contact to large stations 1.0
increased over time (interaction F6,144 = 2.241, p \ 0.043;
station size F2,48 = 5.896, p = 0.005; time F3,144 = 2.376, 0.5
p = 0.072).
0
C
Continuous contacts over time 1.5

The mean number of stations with continuous contact by 1.0

M. gilvus mimicked the pattern observed for total number
of contacts, but more extreme. The number of continuous
0.5
contacts to small stations decreased from a mean of 0.35
stations/mound at week two to zero at week eight. The
0
numbers for medium stations rose slightly from 0.29 to
2 4 8
0.41 stations/mound, whereas those for large stations rose Week
from 0.35 to 1.00 stations/mound over the experimental
period (Fig. 3B). These combined differences were sig- Fig. 3 Mean (±SE) number of stations newly contacted (a),
nificant: continuous contact to large stations increased over continuously contacted (b) and stations abandoned (c) by Macroter-
mes gilvus over the experiment. White circles small stations, grey
time (interaction F6,144 = 6.369, p \ 0.001; station size triangles medium stations, black squares large stations
F2,48 = 4.715, p = 0.014; time F3, 144 = 1.880, p =
0.158).
Abandoned stations

The mean number of stations abandoned by M. gilvus did

not show obvious patterns. The number of abandonments
2.5 to small stations fluctuated around a mean of 0.29 sta-
tions/mound, the mean for medium stations fluctuated
2.0 around 0.22 stations/mound, and the mean for large sta-
Number of staons

tions fluctuated around 0.16 stations/mound over the

experimental period (Fig. 3c). These combined differences
1.5
were significant: fewer large stations were abandoned over
time than small stations (interaction F6,144 = 3.065,
1.0 p = 0.020; station size F2,48 = 0.919, p = 0.406; time
F3,144 = 1.159, p = 0.318).
0.5
Macrotermes carbonarius
0
1 2 4 8 Total contacts to stations
Week
The mean number of contacts by M. carbonarius to stations
Fig. 2 Mean (±SE) total number of stations in contact by Macroter-
mes gilvus over the 8 weeks of the experiment. White circles small
varied between station size and time in a similar pattern as
stations, grey triangles medium stations, black squares large stations M. gilvus, however at lower rates overall. The mean

123
J Pest Sci

number of contacts to small stations was 0.12 sta- A

1.5
tions/mounds after one week, and this rate did not change
over the 8 weeks of the experiment. The mean number of
contacts to medium stations was 0.62 stations/mound after 1.0

1 week, which declined slightly over the remaining

7 weeks to 0.50 stations/mounds. The mean number of 0.5
contacts to large size stations was 0.50 stations/mound after
1 week, rising to 0.75 stations/mound by week eight 0
(Fig. 4). These differences were not significant (interaction B
F6,63 = 1.260, p = 0.289; station size F2,21 = 3.025, 1.5

Number of staons
p = 0.070; time F3,63 = 0.480, p = 0.697).
1.0
New contacts over time
0.5
The mean number of new contacts to stations by M. car-
bonarius mimicked the total number of contacts. The 0
number of new contacts to small stations fluctuated around
C
the mean of 0.13 stations/mound over the 8 weeks of the 1.5
experiment. The number was slightly higher for medium
stations: 0.21 stations/mound, and slightly higher again for 1.0
large stations 0.25 over the experimental period (Fig. 5a).
These differences were not significant (interaction 0.5
F6,63 = 1.474, p \ 0.202; station size F2,21 = 2.100,
p = 0.147; time F3,63 = 1.842, p = 0.149).
0
2 4 8
Continuous contacts over time Week

The mean number of stations with continuous contact by
M. carbonarius showed a different pattern. The number of
continuous contacts to small stations decreased from a
mean of 0.25 stations/mound at week two to zero at week
eight. The numbers for medium stations remained at 0.25
stations/mound, whereas those for large stations rose from
0.38 to 0.75 stations/mound over the experimental period
2.5
2.0
Number of staons
Fig. 5 Mean (±SE) number of stations newly contacted (a),
continuously contacted (b) and stations abandoned (c) by Macroter-
mes carbonarius over the experiment. White circles small stations,
grey triangles medium stations, black squares large stations
(Fig. 5b). These combined differences were significant:
continuous contact to large stations increased over time
(interaction F6,63 = 4.308, p \ 0.005; station size
F2,21 = 1.087, p = 0.355; time F3,63 = 0.538, p = 0.588).
Abandoned stations
The mean number of stations abandoned by M. carbonar-

ius did not show obvious patterns. The number of aban-

1.5 donments of small stations fluctuated around a mean of
0.13 stations/mound, the mean for medium stations fluc-
1.0 tuated around 0.25 stations/mound, and the mean for large
stations fluctuated around 0.21 stations/mound over the
0.5 experimental period (Fig. 5c). These differences were not
significant (interaction F6,63 = 0.722, p = 0.582; station
0 size F2,21 = 0.462, p = 0.636; time F3,63 = 0.072,
1 2 4 8 p = 0.930).
Week
Other termites
Fig. 4 Mean (±SE) total number of stations in contact by Macroter-
mes carbonarius over the 8 weeks of the experiment. White
circles small stations, grey triangles medium stations, black squar-
There were contacts to bait stations by non-target termite
es large stations species. These included species of Coptotermes,

123

Globitermes, Microcerotermes, and Odontotermes. The
mean number of contacts by other termite species, con-
sidered together, to stations varied between station size and
time in a similar pattern as for the Macrotermes species
however at lower rates. Around M. gilvus mounds, the
mean number of contacts to small stations was 0.01 sta-
tions/mounds over the 8 weeks of the experiment; the rate
for medium stations was 0.06 stations/mound, and that for
large size station was 0.22 stations/mound (Fig. 6a). These
combined differences were significant: more large stations
were occupied over time with no change for small sta-
tions (interaction F6,144 = 2.039, p = 0.064; station size
F2,48 = 4.289, p = 0.019; time F3,144 = 1.398, p =
0.246).
The contact rates around M. carbonarius mounds were
higher and also more variable; the mean number of con-
tacts to small stations was 0.28 stations/mounds over the
8 weeks of the experiment; the rate for medium stations
was 0.22 stations/mound, and that for large size stations
was 0.59 stations/mound (Fig. 6b). These differences were
not significant (interaction F6,63 = 0.813, p = 0.564; sta-
tion size F2,21 = 1.820, p = 0.187; effect of time
F3,63 = 0.241, p = 0.867.
Leaf litter
The depth of leaf litter around the mounds varied from 0 to
17.8 cm for M. gilvus, and from 0 to 12.0 cm for M. car-
bonarius. The mean depths were 5.6 ± 0.6 for M. gilvus
1.5 A
1.0
Number of staons
0.5
0 The decreased rate of abandonment in larger stations is
1.5 B
1.0
0.5
0
1 2 4 8
Week
Fig. 6 Mean (±SE) total number of stations contacted by other
termites around Macrotermes gilvus mounds (a) and Macrotermes
carbonarius mounds (b) over the 8 weeks of the experiment. White
circles small stations, grey triangles medium stations, black squar-
es large stations
123
J Pest Sci
and 3.2 ± 0.5 for M. carbonarius; this difference was
significant (t21 = 3.243, p = 0.004).
Discussion
Macrotermes gilvus contacted and occupied large stations
at a much greater rate than the medium baits, and it
eventually shunned small baits. By the end of the experi-
ment, around 50 % of the large stations were occupied by
M. gilvus, but only 5 % of the small baits. M. carbonarius
had a similar pattern between stations of different size,
however the overall contact and occupation rate was lower.
The total rate is a consequence of the discovery of new
stations, and the continued occupation of (and thus the lack
of abandonment of) previously discovered stations. These
were all clearly true of M. gilvus, and to lesser extent of M.
carbonarius, at large stations.
Previous studies have found that larger station size
increased termite contact, or reduced abandonment, plus
greater bait consumption. This pattern was found for two
fungus-growing termites in recent studies. Odontotermes
wallonensis found large wooden blocks faster and infested
them in greater numbers in forests in India (Shanbhag and
Sundararaj 2014). Microtermes mycophagus, found and
infested larger bait stations preferentially in Pakistan; with
80 % of large stations (similar in size to those of the cur-
rent study) infested, which was over three times the rate of
small stations (Iqbal 2014). Non-fungus-growing termites
display the same patterns: including Coptotermes in forests
in Australia (Evans and Gleeson 2006), and various ter-
mites (mostly species in the Apicotermitinae and Nasu-
titermitinae) in forests in Brazil (De Souza et al. 2009). It is
possible that large stations are easier for the termites to
find, or that larger stations are more desirable relative to
other available food resources (Forschler and Ryder 1996;
Perrott et al. 2004), or both.
likely a consequence of lower relative disturbance due to
inspections in the larger stations. Although inspections
were carried out in the same fashion and for the same
period of time for all stations, the relative quantity of wood
disturbed varied according to station size. At each
inspection, just one piece of wood was moved to search for
termites. However, this one piece of wood represented
100 % of the wood in the smallest stations, 50 % of the
wood in the medium stations, and 25 % of the wood in the
large stations. Our results agree with previous studies, that
stations were abandoned with increased frequency of dis-
turbance (Lai 1977; Jones 1990; French et al. 1995; Evans
and Gleeson 2006).
Station size may have unwittingly played a role in
previous baiting studies on fungus-growing termites.
J Pest Sci
Monitoring and baiting stations have been similar in size to
our small stations (Huang et al. 2006) or medium stations
(Peters and Broadbent 2005; Wang et al. 2007; Dhang
2011). These studies reported low total contact rates,
especially for species that do not build mounds (e.g.
8.5 ± 1.9 % in Huang et al. 2006), similar to those
reported in this study; bait stations targeted around
swarming exit holes were more successful (26.0 ± 1.0 %
in Wang et al. 2007). The low contact rates were likely
exacerbated due to the small quantity of bait matrix placed
inside them; the bait matrix was typically completely
removed by termites before each inspection. This neces-
sitated frequent replacement of bait matrix, and this caused
higher disturbance. Placing a quantity of bait sufficiently
large enough to eliminate a colony, once only, is a better
strategy to avoid termites abandoning bait stations (Waller
and La Fage 1987; Lenz 1994; Hedlund and Henderson
1999; Cornelius and Osbrink 2001; Lenz et al. 2012; Evans
and Gleeson 2006).
The results revealed that M. gilvus is a wood feeder, but
M. carbonarius was not. This was not an expected result, as
previous descriptions of both species suggested that both
species were capable of feeding on wood and leaf litter
(Roonwal 1970; Cowie et al. 1989; Lee 2002). In fact, only
M. gilvus was observed to eat the wood in the stations,
whereas the few stations occupied by M. carbonarius
appeared to function as stores for soil dumping, presum-
ably from excavating tunnels. Instead, the lower amount of
leaf litter around M. carbonarius mounds may indicate
their preferred diet. This they collect by foraging above-
ground, in the open air, up to 23 m from their mounds
(Inoue et al. 2001; Hu et al. 2012). In comparison, M.
gilvus has been reported to forage in underground galleries,
or aboveground in protected mud tubes (Acda 2004).
The presence of other species of termites in the bait
stations appeared to be a consequence of the absence of
Macrotermes spp. These other termite species were more
common in larger stations, suggesting that these species,
mostly Coptotermes and Odontotermes, preferred larger
food resources also. These other termite species were more
common in stations around M. carbonarius than M. gilvus
mounds, likely because M. carbonarius had lower infesta-
tion rates, as it did not eat the wood, suggesting that greater
availability was a factor. Finally, the few stations aban-
doned by M. carbonarius were subsequently infested by
other termites, whereas M. gilvus forced other termite spe-
cies to leave stations, reflecting their dominance (Bignell
and Eggleton 2000; Davies et al. 2003; Schuurman 2006).
The purpose of a station is to contain a toxic bait, which
can then be deployed in pest control systems. Termite
control through baiting maybe accelerated if (i) baits are
discovered quickly, (ii) greater numbers of workers are
recruited to the bait stations, so that they (iii) collect and
return greater quantities of bait to the colony. Thus, the
next goal is to find a useful bait toxicant, as the most
commonly used toxicants, chitin synthesis inhibitors, do
not appear to work well with fungus-growing termites,
especially in larger colonies (Lee et al. 2007, 2014). Very
low doses of neurotoxins (e.g. avermectins, fipronil, imi-
dacloprid) or entomopathogenic fungi may be effective
alternative toxicants to chitin synthesis inhibitors to control
infestations of fungus-growing termites (Singha et al. 2010;
Evans and Iqbal 2015; Chen et al. 2015). Fipronil was used
with variable effectiveness in baits against two fungus-
growing termites in Pakistan and China, suggesting more
research on doses rates is needed (Huang et al. 2006; Iqbal
et al. 2015).
We determined in this study that a large size works best
in Macrotermes species, perhaps 10 L or larger for quick
station discovery, and allows recruitment of large numbers
of workers. Once a useful toxicant is found, it could be
used with large stations to develop an effective baiting
system for use against fungus-growing termites, perhaps
other higher termite species as well, in tropical latitudes. If
so, such a baiting system may be deployed not only in
urban areas, but in forestry and agricultural lands as well,
which would allow a relatively environmentally benign
control method to spread to much large areas of the world.
Author contribution statement
NI and TE conceived and designed the study. NI and LSW
conducted the experiments, TE analysed data. NI, LSW,
and TE wrote the manuscript.
Acknowledgments The authors wish to thank the Singapore National
Parks and Singapore Botanical Gardens for granting permission and
providing manpower for the study. NI was supported by the Higher
Education Commission, Pakistan.
Conflict of interest The authors declare no potential conflicts of
interest.
Ethical Approval (Research involving human participants and/or
animals) This research involved neither human nor animal partici-
pation, therefore did not require any ethical approval.
Informed Consent As this research did not involve human partici-
pation, no informed consent was required.
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