Ins. Soc. 42: 359-369 (1995) 1015-1621/95/040359-11 $1.50 + 0.

20/0
9 1995 Birkh~iuser Verlag, Basel

Research article

Foraging activity of the processional termite

Hospitalitermes hospitalis (Termitidae: Nasutitermitinae)
in the rain forest of Brunei, north-west Borneo
D. T. J o n e s ~* and E G a t h o r n e - H a r d y 2
1 Biology Department, Universiti Brunei Darussalam, Bandar Seri Begawan 3186,
Brunei Darussalam
2 Hall Farm, Great Glemham, Saxmundham, Suffolk IP172LP,, UK

Key words: Foraging, Hospitalitermes, termite columns, trails, lichens.

Summary
Four colonies of the column-forming termite Hospitalitermes hospitalis (Haviland) were studied in
a primary forest in Brunei Darussalam, north-west Borneo, and their trails mapped. The termites
leave their nest in the afternoon to graze throughout the night, mainly on lichens growing high up
on the trunk of a canopy tree. Colonies foraged from 46 % to 72 % of nights, and the mean and
maximum trail length was 28.8 m and 65.2 m respectively. Each colony utilized between 14 and 26
trees, with many trees revisited and trails frequently re-used. The mean diameter of the utilized
trees was significantly larger than the mean diameter of trees in the population at the study site.
Termites often made consecutive foraging visits to the largest trees, particularly Shorea spp.
At three colonies per hectare the nest density is lower than expected as more than 90% of
potentially suitable trees are not exploited.

Introduction

T h e majority of termite species travel t h r o u g h tunnels and galleries in soil or w o o d ,

and u n d e r c o v e r e d r u n w a y s over exterior surfaces in search of f o o d (Wood, 1978).
H o w e v e r , in Southeast Asia three g e n e r a of Nasutitermitinae forage a b o v e g r o u n d
in e x p o s e d processional columns; Hospitalitermes, Longipeditermes and Lacessiti-
terrnes. All species are heavily p i g m e n t e d and have relatiely long legs allowing t h e m
free-ranging habits. Species of the genus Hospitalitermes are unusual a m o n g s t
termites in that they feed almost exclusively on free-living, non-vascular plant tissue
which they graze f r o m the surface of three trunks, usually high up in the forest
c a n o p y (Collins, 1988), and which the w o r k e r s carry b a c k to the nest in the f o r m of

* Present address: Biodiversity Division, Department of Entomology, Natural History Museum,

Cromwell Road, London SW7 5BD, UK
360 Jones and Gathorne-Hardy

balls held in the mandibles. In a study of H. umbrinus (Haviland), Collins (1979)

calculated that a foraging column involved up to 500 000 individiual termites from
the colony, and estimated daily consumption at 8.3 mg (dry weight) of food per g
(wet weight) of termites. Examination of the food balls revealed that the material
consisted of crustose lichens and their spores, wood constituents, bryophytes, blue-
green algae and fungal hyphae (Collins, 1979).
Termites are guided along their established routes to feeding areas by following
previously laid odour trails which contain a pheromone produced by the sternal
gland (Stuart, 1969). In addition, Petch (1913) found that H. monoceros (Koenig)
mark their trails with minute specks of proctodeal fluid. Jander and Daumer (1974)
have shown that when exploring new areas, soldier scouts of H. sharpi (which Tho
(1992) regards as conspecific with H. umbrinus) proceed by gravity orientation and
by following crest-lines (i.e. the uppermost surface or ridge-top of elongated objects
such as fallen tree trunks and sticks). Much of the information on this genus is
based on casual observations or qualitative studies (reviewed by Kalshoven, 1958).
A detailed understanding of the factors governing the foraging activities of
Hospitalitermes spp. is still lacking.
Hospitalitermes hospitaIis (Haviland) is a common and conspicuous member of
the termite fauna in the forests of the Belalong area in Brunei Darussalam, north-
west Borneo. If undisturbed, columns of termites leave their nests in the late
afternoon to forage throughout the night and return the following morning. On
some days the last foragers are not back inside the nest until noon. For much of the
night the column consists of two-way traffic; outgoing workers from the nest and
workers returning from the foraging site with balls of food, while soldiers stand
guard on the flanks. The nest sites are variable, ranging from the interior of trunks
or between the roots of living trees, to inside rotten tree stumps or epigeal nests
built over old stumps. Faecal material is usually plastered over the host tree and
forms the nest entrances at which soldiers are constantly present. Our study
monitored the temporal and spatial foraging activities of H. hospitalis and investi-
gated factors which may determine their foraging patterns.

Methods

Study site

Observations were made at Universiti Brunei Darussalam's Plot 1, a permanent

research plot one hectare in size (100 m x 100 m). The plot was located in primary
lowland mixed dipterocarp forest on the east ridge of the Belalong Valley in the
Batu Apoi Forest Reserve (115~ 4~ at an altitude of approximately
285 m. All trees within the plot with a diameter at breast height (dbh) greater than
10 cm had been numbered and many identified where possible. The vegetation of
the Belalong area has been described by Poulsen et al. (in press) and Poulsen (in
press). Preliminary results of studies of the termite fauna in this forest are given by
Jones (in press). Total daily rainfall was recorded at the Kuala Belalong Field
Studies Centre, altitude 60 m, located approximately 600 m horizontal distance
from Plot 1.
Foraging activity of Hospitalitermes 361

Observations
The hectare plot was searched to locate all nests of Hospitaliterrnes spp. The three
nests in Plot 1 and a fourth nest approximately 15 m outside the plot were studied.
Observations were made on 65 days between 01-xi-91 and 29-i-92. At 08.00 h the
four nests were examined and any termite columns present were marked by laying
string alongside their route, from the nest entrance to the point where the column
ascended a tree to forage (here referred to as the "target" tree). When not in use,
the trails were surveyed using tape measure and compass, from the nest entrance to
the target tree. On some occasions a trial could not be followed to its destination so
the length of that trail and its target tree are "unkonwn". Taxonomic identification
of all known targets was not possible, especially outside the plot. The formation of
outgoing columns and their trail utilization were observed on 26 evenings.

Results

Colony density, foraging frequency and trail length

The location of the four colonies and their trails to known target trees are shown in
Fig. 1. The three colonies of H. hospitalis recorded in Plot 1 was the same density as
that found in a hectare plot on the west ridge of the Belalong valley (Jones, in
press), and also for H. umbrinus in Sarawak (Collins, 1979).
It was assumed that a returning column recorded in the morning represented a
successful night of foraging for that colony. Absence of a returning column was
taken to indicate that no foraging had occurred the previous night or that the for-
aging effort had been aborted. The percentage of nights spent foraging (Table 1)
varied from 46 % (colony D) to 72 % (colony A). The maximum number of con-
secutive nights of successful foraging was nine, recorded for colony A (Table 1),
whereas the maximum number of consecutive nights without foraging was eight
(colony B).
The range and mean length of trails to known target trees of each colony is given
in Table 1. Colony C displayed both the maximum trail length (65.2 m) and the mini-
mum (0 m, when the termite column climbed the host tree in which the nest was
located). The actual distance covered by each column was considerably greater than
that reported in Table 1, as the measured trail length excluded the height to which
the column climbed on the target trunk. There was no significant difference in the
trail length between colonies (one way A N O V A ; F = 0.063, df = 3 and 56, p > 0.05).
The mean trail length of the four colonies of H. hospitalis was 28.8 m (SD: + 16.6 m),
which is in line with the estimates of 40 m trail lengths reported for H. umbrinus by
Collins (1979) and Abe (1979), although Jander and D a u m e r (1974) measured one
trail in excess of 300 m.

Utilization of target trees

Table i lists the number of known and unknown targets visited by each colony.
For the period of the study the cumulative total of different targets appears to be
362 Jones and G a t h o r n e - H a r d y

0 50 100
100 ~176 9 * 9 9

Colony A 9 ~
9 ~ ~ 9 9 9

"$ 9 9 9
9 9 -, 9 9 ,*

" 9 ,9 9 9

9 ~
9 ~

9 9 ,9 , ', ~"
Key: ~ ~

99 9 ,. ,9 9
9 Tree over 10 cm dbh 99 9, ;

o Target tree 9 ~ *

9 Nest
: " l ~ , .. .. ;.
Termite trail 50 9: 9 : ;~ 9 .

North
< 9 9
.. ...:..:::-.' ... .. - .. 9 9 Oo,on

Figure 1. Plan of Plot 1 (100 m x 100 m) showing the position of trees within the plot over 10 cm diameter at
breast height, the location of the four H. hospitalis nests, and the foraging trails to known target trees. The arrow
indicates the tree utilized by two colonies

Table 1. Foraging column activity of four coIonies of H. hospitalis located on the east ridge of the Belalong
valley (see Fig. 1), in the Batu Apoi Forest Reserve

Colony A Colony B Colony C Colony D

No. of mornings recorded 65 65 63 61

No. of nights spent foraging 47 (72%) 32 (49%) 30 (48%) 28 (46%)
No. of known target trees 18 11 13 18
No. of unknown target trees 4 1 1 6
No. of active nights foraging 11 (23%) 5 (16%) 1 (3%) 9 (32%)
at more than one target
Maximum n u m b e r of consecutive 9 5 3 2
nights of foraging
Maximum n u m b e r of consecutive 5 3 2 2
nights at same target
Maximum n u m b e r of consecutive 3 8 2 5
nights without successful foraging
Minimum trail length (m) 3.6 4.4 0.0 2.4
Maximum trail length (m) 55.3 53.1 65.2 53.3
M e a n trail length (m) (_+1 SD) 27.9 (18.0) 27.5 (15.2) 30.2 (18.8) 30.0 (14.6)
Foraging activity of Hospitalitermes 363

Colony D
25-
= i l i a
Colony A
= n u n n n 9 1 4 9

=== a B a g a i

20 84 9 J = m = = =

JJ = = = =

I = n n

===

===

15 Colony C
=== o o o o o 0
o o o o o o
G)
9 0 0 0 e e l l e a Colony B
9 0 0 0 e e e o * *

10 9 oo = , o o o e

=oo o e 9
o
oo 9 1 4 9 1 4 9

9 =e
O * =

O=

. . . . . . . . . i . . . . . . . . . n . . . . . . . . . n . . . . . . . . . n . . . . . . . . . i . . . . . . . . . n
0 10 20 30 40 50 60

Number of nights foraging

]Figure 2. The cumulative number of known and unknown target trees utilized by each colony of H. hospitalis
during the study period. Each unknown target was considered to be a "new" tree

approaching an upper limit for each colony (Fig. 2). The number of trees utilized,
estimated from Fig. 2, ranged from about 14 (colony B) to about 26 (colony D).
Of the 592 trees in Plot 1 with a dbh greater than 10 cm, only 23 (3.9 %) were exploit-
ed by Hospitaliterrnes during the study period. Using the estimates of the maximum
number of targets for colonies A, B and C (total 53 trees), even if all these targets
had been inside the plot, H. hospitalis would still be exploiting less than 10% of
trees over 10 cm dbh per hectare. Colony D had no targets inside Plot i (see Fig. 1)
and other neighbouring colonies were not seen foraging within the plot.
Each colony's foraging range was usually separate from the other colonies' but
at one location two did meet. Colonies B and C both exploited the same individual
tree (see Fig. 1), a Shorea sp. of 148.8 cm dbh, the largest tree in the plot. A column
from both colonies was never recorded on this tree on the same night, and the trail
from each nest went up opposite sides of the trunk. However, it is not known if, and
how, colonies interact to maintain separate feeding ranges.
The dbh of known target trees ranged from 8.0 cm to 148.8 cm, with only two
targets less than 10 cm dbh. The mean dbh (_+SD) of known targets was 52.9 cm
(+_34.3 cm), whereas it was 22.7 cm (_+17.5 cm) for all trees in the plot with a dbh
greater than 10 cm. As the distribution of trees is heavily skewed in favour of
smaller diameters a non-parametric test was performed to ascertain if there was a
difference between target and non-target trees. Target trees (excluding the two
targets with a dbh less than 10 cm) had a significantly larger dbh than the non-target
364 Jones and Gathorne-Hardy

trees in the plot (Mann-Whitney U test= 1732.5, p <0.001). Combining the data
from all colonies there was a significant positive correlation between the dbh of
target trees and the number of recorded visits to each tree (on a double log10 scale
r 2= 0.284, p > 0.05). There was no significant correlation between the trail length and
the log10 dbh of each target (r 2= 0.049, p > 0.05), nor was there a significant correla-
tion between the trail length and the number of visits to each target (r 2=0.031,
p > 0.05).
The known target trees span a wide range of taxa, representing eleven families
(Table 2). Shorea spp. were the most common target (31% of identified targets)
with eight individual trees receiving 39 visits from foraging columns during the
study period. All other taxa were represented by three or less targets. There was a
higher than expected number of Shorea spp. in the target population when compar-
ed to the number of individual trees of Shorea spp. in the population of Plot 1 with
a dbh greater than 10 cm (employing Yate's correction: X2= 6.35, df= 1, p < 0.05).
Shorea is one of the predominant genera amongst the big trees of Borneo's mixed
dipterocarp forest (Whitmore, 1984). Of the nine trees in Plot 1 with a dbh greater
than 80 cm, seven were Shorea spp., while an investigation on the west ridge of the
Belalong valley revealed that 44.3 % of the total basal area of all trees over 10 cm
dbh in a hectare plot was contributed by 76 individuals of eighteen Shorea species
(Poulsen et al., in press). Therefore, H. hospitalis may simply be selecting larger
trees to forage upon rather than being attracted to Shorea species per se. However,
the possibility that Shorea bear more of the favoured lichen compared with other
trees of similar size cannot be ruled out.
All four colonies show some degree of consecutive foraging on particular
targets. This was most pronounced in colony A (Fig. 3) which visited the same tree
(138.8 cm dbh) nine times between days 16 to 28 of monitoring, and visited another
target (121.0 cm dbh) six times between days 53 and 63. The simultaneous exploita-
tion of two or more target trees by the same colony was a common phenomenon.
The highest incidence of multiple targets, on 32 % of active nights, was recorded for
colony D (Table 1). Often a colony maintained two (occasionally three) columns on
independent trails, each to a different target. In other cases the first target was en
route and some of the column members continued along the trail to exploit a second
target, whilst sometimes a single column proceeded along a trail from the nest and
then split to follow two separate trails (see Fig. 1).

Column formation and m o v e m e n t

Column formation could start any time after mid afternoon but generally
commenced in the late afternoon. Termites usually emerged from several nest
exists and numbers outside would quickly build up, frequently resulting in the
formation of several columns. After a period of disorganized activity (occasionally
lasting up to six hours) during which time the different columns would indepen-
dently advance short distances and retreat, one or two columns would become
firmly established and the remaining columns would retire into the nest. Advance
scouts were not observed and individuals never moved great distances beyond the
head of the developing column.
F o r a g i n g activity of Hospitalitermes 365

Table 2. Generic or family level identity of the known target trees, the number of visits made to those trees
during the study period by foraging columns from colonies A, B and C of H. hospitalis, and the percentage of
the tree population (diameter at breast height > 10 cm) in the plot represented by each taxon. The targets of
colony D were not identified

G e n u s / F a m i l y of t a r g e t tree No. of visits No. of t r e e s visited % of p o p u l a t i o n

in h e c t a r e plot

Shorea 39 8 11.0
Tristania 12 2 0.3
Cratoxylon 9 2 0.3
Eugenia 7 3 6.4
Canarium 5 1 ?
Elateriospermum 5 2 3.2
Knema 2 1 ?
Lauraceae 2 1 4.2
Vatica 2 1 3.2
Tiliaceae 2 1 0.5
Sindora 1 1 ?
Dialium 1 1 0.3
Anacardiaceae 1 1 1.0
Scaphium 1 1 0.5
Memecylon 1 1 0.7
Nephelium 1 1 1.2
Indeterminate 35 ?

Total 126

I I
~,C 15
I 1
-gE
~2 I I
.--. 7: I I
m
I I
I I
I I
I I
" I I

1
Tree diameter:
9 dbh < 2 0 cm I I
9 dbh 2 0 - 4 0 cm I I
9 dbh 4 0 - 8 0 cm
9 dbh > 80 cm
o dbh unknown
0 .II.. 10
9 i!!
20
I
30
]
40 50
I
,
I
,
60
[

Days on which nest was monitored for foraging activity

Figure 3. The occurrence of foraging activity by H. hospitalis (colony A) and the utilization of trees ranked in
order of increasing trail length. Vertical lines indicate successful foraging on one or more trees. Dashed lines
indicate unknown trail length. Tree diameter at breast height is indicated by the size of the circle
366 Jones and Gathorne-Hardy

Many trails shared the initial part of their route leading from the nest before
branching off to different targets. For the greater part of the trail the column of
termites remained off the forest floor, usually marching along stems, vines, roots
and fallen branches. Columns revisiting a target usually followed the same trail
almost exactly, with only minor changes in the route due possibly to damage or
disturbance to part of the trail. When in progress, a column would sometimes
divide, only to rejoin again after a short distance. The maximum time a trail remain-
ed unused over a period of continuous monitoring was fifteen days (colony D).
Despite such long intervals between use, the termites were able to follow the origin-
al trail with great precision. Marking the trail with string appeared not to deter its
use by termites.

The influence o f rain on foraging behaviour

If rain occurred during the period of column formation the outgoing termites
retreated to the nest but re-emerged en mass once the rain had stopped. Rain at any
time after the column was established caused the termites to stop marching and
disperse to find shelter alongside the trail. Foraging recommenced once the rain
ceased. The same response to rain is also reported for H. umbrinus and H. mono-
ceros (Collins, 1979; Petch, 1913). This response was similar to that seen during any
physical disturbance to the column.
Differences in foraging frequency between the four colonies made it difficult to
ascertain if heavy or prolonged rainfall caused a complete failure of foraging. For
example, successful foraging was not recorded for any of the four colonies on the
night of the heaviest daily rainfall (106 mm) but on the night of the second heaviest
daily rainfall (72.5 ram) foraging was reported for two of the colonies. However,
colonies were often inactive in the absence of any rainfall, and for one night the four
colonies did not forage when the daily rainfall was only 0.5 mm.

Discussion

In lowland mixed dipterocarp forest in Sarawak, H. umbrinus by-pass easily

accessible mosses, algae and fungi near the nest to make long excursions into
the canopy to collect lichens (Collins, 1979). Only once during our study was
H. hospitalis observed feeding on mosses near ground level. Collins (1983) suggests
that Hospitalitermes spp. show a preference for lichens because they offer an
important dietary source of nitrogen. Large diameter trees, such as Shorea, have
a disproportionately greater surface area than smaller trees and therefore have
the potential to carry a far greater amount of lichens, and indeed H. hospitalis
conduct significantly more visits to trees of larger diameter. The temporal
pattern revealed in Fig. 3 includes bouts of consecutive nights foraging on two
trees of large dbh, and also suggests that targets with a smaller dbh sustain
fewer consecutive nights of foraging. This assumes that the termites forage ex-
clusively on the target and do not move to adjacent trees via contacts between tree
crowns.
Foraging activityof Hospitalitermes 367

The upper limit to the cumulative total of target trees (see Fig. 2) suggests that
the number of targets currently used by each colony may remain approximately
constant over time. However, individual targets would be replaced if sufficiently
depleted of lichens, and as new trails were developed in response to disturbances to
established trails due to events such as tree falls. The location of new target trees
appears to be indiscriminate and is probably the result of the termites advancing in
a random direction as scouts follow the "crest-line" of objects they encounter until
a tree of suitable size is reached. This is reflected in the indirect, often tortuous
route of trails to targets (see Fig. 1). Wood (1978) notes that other termite genera,
for example Nasutiterrnes guayanus (Holmgren) which can travel 50 m under cover-
ed runways to feed, seldom follow the most direct route to a food source. Hospita-
litermes are highly effective in defense while foraging in the open. Soldiers can
squirt sticky, irritating chemicals from their enlarged frontal gland to deter pre-
dators (Chuah et al., 1983; Prestwich, 1984), and Petch (1913) suggested that crest-
lining along narrow objects helps protect the column from ambush by ants.
Considering the limited area over which target trees are distributed, and the
relatively large area of the plot which termite columns do not enter, why is the nest
density only 3 per hectare? As more than 90 % of trees in the plot are not exploited
by H. hospitalis but presumably offer sufficient food resources to support more
colonies, the nest density may be limited by factors other than food availability. One
possibility is that intense interspecific competition for nesting sites means that such
sites are not readily available (Emerson, 1955). Several species of termite, as well as
many ant species and other groups of animals, utilize microhabitats which are
similar to those used as nesting sites by H. hospitalis. If suitable nesting sites are few,
and the alates' powers of dispersal are weak, then the chance of a pair of alates
finding an unoccupied nesting site and successfully establishing a new colony may
be very low.
The only comparable results on Hospitalitermes foraging frequency are those for
H. monoceros. Petch (1913) observed a single colony for 119 consecutive days and
recorded foraging activity on 58 % of nights, which is within the range reported in
our study. However, the maximum consecutive nights of foraging (31 days) and
inactivity (15 days) are longer than those of H. hospitalis (Table 1). Jander and
Daumer (1974) stated that H. umbrinus store food in the nest (but no description
was given) and suggested that foraging activity was triggered by the depletion of
these stores. However, other studies do not provide evidence of any species of
Hospitalitermes storing food. Petch (1913) reported that H. monoceros workers
carry the balls of food about inside the nest for non-foraging members of the colony
to consume.
This study raises many questions concerning the factors which control the choice
of targets and the foraging frequency of Hospitaliterrnes. There is little information
regarding the use of permanent foraging trails by termites but considerably more
about ants (see Brian, 1983). In the case of harvester ants (Pogonomyrrnex spp.),
foragers travel up to 40 m on well established and pheromonally marked "trunk
trails" (H611dobler and Lumsden, 1980) before diverging on individual excursions
where interactions with neighbouring conspecific colonies are most likely to occur.
The decision as to which of its habitual trails P. barbatus will use depends in part on
the encounters of the previous day (Gordon, 1991). To a major extent H. hospitalis
368 Jones and Gathorne-Hardy

is isolated from its conspecific neighbours so the daily selection of targets may
be influenced instead by rates of predation or disturbance on the trails previously
used.
The amount of food collected on a foraging excursion depends to a large extent
on the numbers of workers visiting the feeding site, and this in turn is influenced by
the trail length. For example, in the case of colony C, when the longest (65.2 m) and
the shortest (0 m) trails were used and both columns were maintained for approx-
imately the same duration, considerably more workers could have visited the feed-
ing site on the shortest trail, if the numbers of termites per unit length of column
was the same on both trails. This assumes that H. hospitalis have a constant walking
speed on established trails, which is the case for H. umbrinus (Collins, 1979). How-
ever, this picture is complicated by Collins' results (1979) which demonstrate that
the rate at which H. urnbrinus leave the nest can vary throughout a single foraging
excursion and from night to night. In addition, Jander and D a u m e r (1974) infer
that individual termites of H. urnbrinus make only one journey to the target per
night of foraging, and this was also implicit in Collins' (1979) calculations of for-
aging activity. But on the shortest trails workers of H. hospitalis would have the
opportunity to make more than one journey to the feeding site.
Given differing trial lengths and column densities, single and multiple target util-
ization, and the possibility of workers making more than one visit to the foraging
site in a night, the amount of food collected per night may be highly variable. The
fact that H. hospitalis trail length is not significantly correlated with either target
dbh or the number of visits to each target may suggest that walking is not energet-
ically expensive relative to the value of the food balls collected and thus the colony
can support longer foraging excursions. Fewell (1988) found that the energetic costs
incurred by the seed-harvesting ant Pogonornyrrnex occidentalis while foraging
were very low in relation to the value of the seeds collected, and therefore minimiz-
ing foraging distance was not a major benefit.
When constructing models of social insect foraging (revied by Ydenberg and
Schmid-Hemple, 1994), Hospitaliterrnes offers a more simplified system than
ants because the former do not have the added complexity of workers foraging
individually and recruiting nest mates to food sources. As trails can be mapped, and
the quantity of food brought back to the nest can be estimated (Collins, 1979),
Hospitalitermes is an ideal candidate for further studies into the energetics of
foraging behaviour.

Acknowledgements
We thank the staff of the joint Universiti Brunei Darussalam - Royal Geographical Society Rain
Forest Project 1991/92, under whose auspices and support this study was conducted. This work was
undertaken while DTJ was a Research Fellow at UBD. We are grateful to Mark Collins for his
encouragement, and Gathorne Cranbrook for assisting with field work. Our special thanks go to
Yulis, Wathinah and Ramlah and the rest of the staff at the Kuala Belalong Field Studies Centre
for their friendship and cooperation. Bert Orr, Paul Eggleton, Bill Sands and Nigel Stork very
kindly provided constructive comments on the manuscript.
Foraging activity of Hospitalitermes 369

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Received 21 June 1994;

revised 15 March 1995;
accepted 24 March 1995.