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

Macrotermes carbonarius followed a similar pattern but 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

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 J Pest Sci

various methods to prevent and control termite attack, components of current commercial baits are still being
including physical barriers, chemical insecticide barriers in tested against them (e.g. Lee et al. 2014).
soil and in building materials, dusting with insecticide It seems likely that the major biological and ecological
toxicants, and baiting (Edwards and Mill 1989; Pearce differences between fungus-growing termites and the
1997; Hu 2011). Each method has various advantages rhinotermitid termites could drive different bait station
depending on the building construction type and density of design. Food size and bait station design can have large
buildings, from free-standing wooden houses in low den- difference in rhinotermitid termite foraging behaviour
sity suburbs to high rise concrete apartment blocks in the (Evans and Gleeson 2006; Lenz et al. 2009; Evans 2010).
high density inner city. Each method has differing aims, In this study, we investigated the effect of bait station
from how they preventing the termites from causing design on two Macrotermes species in Singapore. We
damage, to minimizing undesirable side effects, such as considered the effect of size in attracting and retaining
negative environmental impacts. foraging termites, as this is a critical attribute in the success
Baiting was developed to reduce the population size of, of any bait system (Evans and Iqbal 2015), and will need to
or even eliminate termite colonies, i.e. kill all the termites, be optimized to make baiting systems work against fungus-
and to be more environmentally benign. The colony was growing termites in tropical systems. We hypothesized that
eliminated by feeding insecticides in a cellulose food matrix larger bait stations with greater food resources would be
to the termites, and this approach produced the environment found more quickly, recruit more termites, and retain those
benefits: using less insecticide and containing the insecti- termites when disturbed during inspections.
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 Materials and methods
the second most used termite control methods, at least in
wealthier countries (Evans and Iqbal 2015). Study site
Current commercial baiting systems may not be useful
everywhere. Current baiting systems were designed in We performed the study at the Singapore Botanic Gardens
temperate latitudes, especially in the USA, for use in urban (SBG; Latitude 1.3120511N, Longitude 103.817417E).
habitats, and to target pest termites in the Rhinotermitidae, The SBG is 74 hectares and contains a mixture of habitats,
specifically species in Reticulitermes and Coptotermes (Su including primary dipterocarp rainforest, various stages of
1995; Grace et al. 1996). There are many other pest ter- secondary forests, manicured parklands, and specialist
mites found in the tropics, in rural and forested habitats, gardens. Mound-nests of two Macrotermes species,
and in other termite families (Roonwal 1979; Cowie et al. Macrotermes gilvus and Macrotermes carbonarius, are
1989; Mitchell 2002; Constantino 2002). It is unclear found in the forests and parkland, often near large trees.
whether the current commercial baiting systems are We selected a total of 25 mound-nests: 17 M. gilvus and
appropriate for these much wider range of circumstances; eight M. carbonarius. The mounds were all considered to
in tropical Asia they are generally used only in urban contain healthy colonies, with complete clay covering,
habitats against Coptotermes species (Lenz 2002; Lenz and recent construction activity, and termites foraging nearby.
Evans 2002; Lee et al. 2007; Evans and Iqbal 2015). The M. gilvus mounds were significantly smaller than those
One termite subfamily, the Macrotermitinae, may be of of M. carbonarius mounds (t tests; height t = 1.989,
particular concern: various species of Macrotermes, Mi- df = 21, p = 0.059; width t = 3.409, df = 21, p = 0.003;
crotermes, Odontotermes and other genera are important Fig. 1). All these mounds were present near trees, and we
pests in tropical Asia and Africa (Roonwal 1970; Wood observed Macrotermes galleries in several trees, some of
et al. 1980, 1987; Cowie et al. 1989; Tiben et al. 1990). which were culturally important and were under heritage
These termites do not have protozoan symbionts in their protection laws.
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). Bait stations
In general, they have larger body sizes, live in more pop-
ulous colonies, forage over longer distances, dominate We tested three bait/monitoring station sizes. All were
other termites living in their habitats, and are significant constructed from cylindrical plastic containers with sloping
components of these habitats (Evans et al. 1998; Bignell sides, with holes of 8 mm diameter, on sides and base, and
and Eggleton 2000; Davies et al. 2003; Schuurman 2006; with a piece of wood in proportion to the size of the con-
Pringle et al. 2010). Fungus-growing termites may have tainer. The small stations were ca. 0.35 L (16.5 cm high,
different requirements to the Rhinotermitidae; the top [ 8 cm, bottom [ 2.5 cm), and contained one piece of

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250 Data recording and analysis

We inspected all stations after 1, 2, 4, and 8 weeks. At each

Mound Diameter (cm)

200
inspection, we cleared the covering soil, moved the paver,
150 opened the lid, and visually checked for the presence of
termites or their mudding in each station. We attempted to
100
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
50
immediately replace the lid and paver, and cover with soil.
After the first week inspection, we recorded new contact,
0
0 20 40 60 80 100
continuous contact, and abandonment by the respective
Mound height (cm) Macrotermes species and other, non-target, termite species.
We defined ‘new’ contact to a station as the first appear-
Fig. 1 Sizes of Macrotermes gilvus and M carbonarius mounds from ance of that species in that station, from inspection 2
the Singaporean Botanic Gardens used in the study. Black onwards, and ‘continuous’ contact as the same species in
squares = M. gilvus; white circles M. carbonarius. Regressions M.
gilvus y = 0.8207x ? 47.098, r2 = 0.16; M. carbonarius y =
that station as observed in the previous inspection. We
1.6084x ? 62.6, r = 2, R2 = 0.79 defined ‘abandonment’ as the loss of a species in a station
in which it has been observed in the previous inspection.
Pinus radiata wood (12.1 9 5.0 9 2.5 cm). This size was We analysed data using repeated measures ANOVA,
comparable to the earliest commercial systems, including because the units were sampled (i.e. inspected) repeatedly
Sentricon by Dow Chemical Company and Firstline by (Sokal and Rohlf 1995). The ‘repeated’ factor was time,
FMC Corporation. The medium stations were ca 3.6 L in with time interval adjusted for weeks (1, 2, 4 and 8) rather
volume (15.5 cm high, top [ 18.5 cm, bottom [ 16 cm), than inspection number (1, 2, 3, and 4). As each inspection
and contained two pieces of wood (13.0 9 7.6 9 2.5 cm). of each station was not considered independent in the
This size was comparable to later design bait systems, analysis, there was no pseudoreplication. We performed all
including Exterra by Ensystex, Nemesis by PCT analyses in SYSTAT (v. 9.0, 1998).
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 Results
(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. Macrotermes gilvus
Another piece of wood (26.5 9 15.2 9 2.5 cm) was
placed underneath each large station. The large station was Total contacts to stations
comparable in size to non-commercial monitoring stations
used in previous studies (Evans et al. 1998, 1999; Evans The mean number of contacts by M. gilvus to stations
and Gleeson 2001; Evans 2004, 2006; Lenz et al. 2012, varied between station size and time. The mean number of
2013). contacts to small stations was 0.53 stations/mound after
We installed four large, four medium, and four small 1 week, but this eventually decreased to 0.18 after eight
stations per mound for a total of 300 stations around the 25 weeks. The mean number of contacts to medium stations
mounds. We dug holes suitable in size to accommodate the was similar to that of small stations after one week, 0.59
different-sized stations at ca. 2 m from the edge of the stations/mound, which remained steady until eight weeks,
mounds. We positioned the holes around the circle, so that 0.65 stations/mounds. The mean number of contacts to
adjacent holes were 30 to each other, relative to the centre large size stations was similar to the small and medium
of the mound, each hole and station ca. 1.5 m apart. We stations after 1 week, 0.47 stations/mound; however, this
place stations sequentially in size sets around the circle, i.e. increased fourfold to 1.94 stations/mound after 8 weeks
small, medium, large, small, medium, etc. The stations (Fig. 2). These combined differences were significant:
were placed in the holes, covered with plastic lids (small [ fewer small stations were occupied over time, whereas the
12 cm, medium [ 21 cm, large [ 28 cm), on top of which opposite was true for large stations (interaction
we placed concrete pavers and soil to prevent disturbance F6,144 = 10.155, p \ 0.001; station size F2,48 = 7.143,
from members of the public. p = 0.002; time F3,144 = 4.949, p = 0.003).

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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

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

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,

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Globitermes, Microcerotermes, and Odontotermes. The and 3.2 ± 0.5 for M. carbonarius; this difference was
mean number of contacts by other termite species, con- significant (t21 = 3.243, p = 0.004).
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 Discussion
mean number of contacts to small stations was 0.01 sta-
tions/mounds over the 8 weeks of the experiment; the rate Macrotermes gilvus contacted and occupied large stations
for medium stations was 0.06 stations/mound, and that for at a much greater rate than the medium baits, and it
large size station was 0.22 stations/mound (Fig. 6a). These eventually shunned small baits. By the end of the experi-
combined differences were significant: more large stations ment, around 50 % of the large stations were occupied by
were occupied over time with no change for small sta- M. gilvus, but only 5 % of the small baits. M. carbonarius
tions (interaction F6,144 = 2.039, p = 0.064; station size had a similar pattern between stations of different size,
F2,48 = 4.289, p = 0.019; time F3,144 = 1.398, p = however the overall contact and occupation rate was lower.
0.246). The total rate is a consequence of the discovery of new
The contact rates around M. carbonarius mounds were stations, and the continued occupation of (and thus the lack
higher and also more variable; the mean number of con- of abandonment of) previously discovered stations. These
tacts to small stations was 0.28 stations/mounds over the were all clearly true of M. gilvus, and to lesser extent of M.
8 weeks of the experiment; the rate for medium stations carbonarius, at large stations.
was 0.22 stations/mound, and that for large size stations Previous studies have found that larger station size
was 0.59 stations/mound (Fig. 6b). These differences were increased termite contact, or reduced abandonment, plus
not significant (interaction F6,63 = 0.813, p = 0.564; sta- greater bait consumption. This pattern was found for two
tion size F2,21 = 1.820, p = 0.187; effect of time fungus-growing termites in recent studies. Odontotermes
F3,63 = 0.241, p = 0.867. wallonensis found large wooden blocks faster and infested
them in greater numbers in forests in India (Shanbhag and
Leaf litter Sundararaj 2014). Microtermes mycophagus, found and
infested larger bait stations preferentially in Pakistan; with
The depth of leaf litter around the mounds varied from 0 to 80 % of large stations (similar in size to those of the cur-
17.8 cm for M. gilvus, and from 0 to 12.0 cm for M. car- rent study) infested, which was over three times the rate of
bonarius. The mean depths were 5.6 ± 0.6 for M. gilvus 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-
1.5 A
mites (mostly species in the Apicotermitinae and Nasu-
titermitinae) in forests in Brazil (De Souza et al. 2009). It is
1.0 possible that large stations are easier for the termites to
find, or that larger stations are more desirable relative to
Number of staons

0.5 other available food resources (Forschler and Ryder 1996;

Perrott et al. 2004), or both.
0 The decreased rate of abandonment in larger stations is
likely a consequence of lower relative disturbance due to
1.5 B
inspections in the larger stations. Although inspections
were carried out in the same fashion and for the same
1.0 period of time for all stations, the relative quantity of wood
disturbed varied according to station size. At each
0.5 inspection, just one piece of wood was moved to search for
termites. However, this one piece of wood represented
0
100 % of the wood in the smallest stations, 50 % of the
1 2 4 8 wood in the medium stations, and 25 % of the wood in the
Week large stations. Our results agree with previous studies, that
stations were abandoned with increased frequency of dis-
Fig. 6 Mean (±SE) total number of stations contacted by other turbance (Lai 1977; Jones 1990; French et al. 1995; Evans
termites around Macrotermes gilvus mounds (a) and Macrotermes
and Gleeson 2006).
carbonarius mounds (b) over the 8 weeks of the experiment. White
circles small stations, grey triangles medium stations, black squar- Station size may have unwittingly played a role in
es large stations previous baiting studies on fungus-growing termites.

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Monitoring and baiting stations have been similar in size to return greater quantities of bait to the colony. Thus, the
our small stations (Huang et al. 2006) or medium stations next goal is to find a useful bait toxicant, as the most
(Peters and Broadbent 2005; Wang et al. 2007; Dhang commonly used toxicants, chitin synthesis inhibitors, do
2011). These studies reported low total contact rates, not appear to work well with fungus-growing termites,
especially for species that do not build mounds (e.g. especially in larger colonies (Lee et al. 2007, 2014). Very
8.5 ± 1.9 % in Huang et al. 2006), similar to those low doses of neurotoxins (e.g. avermectins, fipronil, imi-
reported in this study; bait stations targeted around dacloprid) or entomopathogenic fungi may be effective
swarming exit holes were more successful (26.0 ± 1.0 % alternative toxicants to chitin synthesis inhibitors to control
in Wang et al. 2007). The low contact rates were likely infestations of fungus-growing termites (Singha et al. 2010;
exacerbated due to the small quantity of bait matrix placed Evans and Iqbal 2015; Chen et al. 2015). Fipronil was used
inside them; the bait matrix was typically completely with variable effectiveness in baits against two fungus-
removed by termites before each inspection. This neces- growing termites in Pakistan and China, suggesting more
sitated frequent replacement of bait matrix, and this caused research on doses rates is needed (Huang et al. 2006; Iqbal
higher disturbance. Placing a quantity of bait sufficiently et al. 2015).
large enough to eliminate a colony, once only, is a better We determined in this study that a large size works best
strategy to avoid termites abandoning bait stations (Waller in Macrotermes species, perhaps 10 L or larger for quick
and La Fage 1987; Lenz 1994; Hedlund and Henderson station discovery, and allows recruitment of large numbers
1999; Cornelius and Osbrink 2001; Lenz et al. 2012; Evans of workers. Once a useful toxicant is found, it could be
and Gleeson 2006). used with large stations to develop an effective baiting
The results revealed that M. gilvus is a wood feeder, but system for use against fungus-growing termites, perhaps
M. carbonarius was not. This was not an expected result, as other higher termite species as well, in tropical latitudes. If
previous descriptions of both species suggested that both so, such a baiting system may be deployed not only in
species were capable of feeding on wood and leaf litter urban areas, but in forestry and agricultural lands as well,
(Roonwal 1970; Cowie et al. 1989; Lee 2002). In fact, only which would allow a relatively environmentally benign
M. gilvus was observed to eat the wood in the stations, control method to spread to much large areas of the world.
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 Author contribution statement
leaf litter around M. carbonarius mounds may indicate
their preferred diet. This they collect by foraging above- NI and TE conceived and designed the study. NI and LSW
ground, in the open air, up to 23 m from their mounds conducted the experiments, TE analysed data. NI, LSW,
(Inoue et al. 2001; Hu et al. 2012). In comparison, M. and TE wrote the manuscript.
gilvus has been reported to forage in underground galleries,
or aboveground in protected mud tubes (Acda 2004). Acknowledgments The authors wish to thank the Singapore National
Parks and Singapore Botanical Gardens for granting permission and
The presence of other species of termites in the bait providing manpower for the study. NI was supported by the Higher
stations appeared to be a consequence of the absence of Education Commission, Pakistan.
Macrotermes spp. These other termite species were more
common in larger stations, suggesting that these species, Conflict of interest The authors declare no potential conflicts of
interest.
mostly Coptotermes and Odontotermes, preferred larger
food resources also. These other termite species were more Ethical Approval (Research involving human participants and/or
common in stations around M. carbonarius than M. gilvus animals) This research involved neither human nor animal partici-
mounds, likely because M. carbonarius had lower infesta- pation, therefore did not require any ethical approval.
tion rates, as it did not eat the wood, suggesting that greater Informed Consent As this research did not involve human partici-
availability was a factor. Finally, the few stations aban- pation, no informed consent was required.
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 References
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