Crop Protection 22 (2003) 87–93

Effects of maize–legume intercrops on termite damage to maize,

activity of predatory ants and maize yields in Uganda
B.M. Sekamattea,*, M. Ogenga-Latigob, A. Russell-Smithc
a
Namulonge Research Institute, P.O. Box 7084, Kampala, Uganda
b
Department of Crop Science, Faculty of Agriculture, Makerere University, P.O. Box 7062, Kampala, Uganda
c
Natural Resources Institute, University of Greenwich, Central Avenue, Chatham Maritime, Kent ME4 4TB, UK
Received 21 May 2001; accepted 24 June 2002

Abstract

Field experiments during two cropping seasons at Namulonge Agricultural Research Institute, Uganda assessed the effect of
intercropping maize with soyabean, groundnut and common beans on termite damage to maize, activity of common predatory ants
and maize yields. Intercropping caused a significant (Po0:01) reduction in termite attack, reduced loss in grain yield of maize and
increased the nesting of predatory ants in maize fields. In many instances, termite attack was significantly lower (P ¼ 0:05) in the
maize–soyabean intercrop than in maize intercrops with groundnuts and beans. Species of the genera Myrmicaria and Lepisiota were
the dominant ant predators recorded. The study revealed that soyabean and groundnut are more effective in suppressing termite
attack than common beans, suggesting the necessity to identify suitable legumes for each cropping situation. Overall, intercropping
might form a component of an integrated management strategy for termites in smallholder cropping systems in East Africa.
r 2002 Elsevier Science Ltd. All rights reserved.

Keywords: Intercropping; Termites; Lepisiota; Myrmicaria; Maize; Uganda

1. Introduction In the tropics, subsistence farmers who practice low

Termites (Isoptera: Termitidae) are one of the most
important maize production constraints in Uganda
(Sekamatte et al., 2001). Feeding damage on roots,
stem bases and leaves frequently results in plant lodging
and damage to cobs causing yield losses between 10%
and 30% in different parts of the country (Sekamatte,
2001). Control of termite pests currently relies mainly on
the use of chemical insecticides (Anon., 1998) but the use
of non-chemical control methods has been emphasised
following the ban on persistent organochlorine insecti-
cides (Logan et al., 1990). Such methods attempt to (i)
prevent termite access to plants, (ii) reduce termite
numbers in the vicinity of plants, and (iii) reduce
susceptibility or increase resistance of the plants
themselves (Logan et al., 1990). Although there is little
published information on non-chemical control of
termites in annual crops, cultural practices such as
intercropping, have been reported to reduce termite
infestation in forestry (Tho, 1974).
*Corresponding author.
input agriculture use intercropping to minimise the risk
of crop failure and to improve the yields of particular
crops (Ofori and Stern, 1987). For example, in Uganda,
over 70% of farmers grow maize as an intercrop
(Sekamatte, 2001). Intercrops have been reported to
reduce pest incidence and damage to the principal crop
(Baliddawa, 1985; Altieri and Letourneau, 1982; Risch
et al., 1983; Trenbath, 1993) and, in Uganda, have been
proposed as a component of IPM strategies for beans
(Kyamanywa and Tukahirwa, 1988; Ogenga-Latigo
et al., 1992). Intercropping of maize with napier grass
and Desmodium sp. in Kenya has been proposed against
lepidopteran stem borers and Striga in maize (Khan
et al., 2000). A recent study (Ekesi et al., 1999) provided
some evidence that intercropping improved the control
of thrips in cowpea by the entomo-pathogenic fungus
Metarhizium anisopliae.
Although little is known about possible preference of
termites for particular crops, farmers’ reports and our
own observations suggested that intercropping might
reduce termite damage to maize. In parts of northern
Uganda, maize intercropped with sorghum usually

0261-2194/02/$ - see front matter r 2002 Elsevier Science Ltd. All rights reserved.
PII: S 0 2 6 1 - 2 1 9 4 ( 0 2 ) 0 0 1 1 5 - 1
88 B.M. Sekamatte et al. / Crop Protection 22 (2003) 87–93
suffers lower termite damage compared to pure stands
(Farmers’ reports.) Farmers in this region regarded the
sorghum plants as reservoirs of the black ant ‘ngini–
ngini’ (Lepisiota sp.), which we have observed to prey on
a range of pests and which are valued by the farmers,
especially on cotton (Sekamatte et al., in press). Such
farmers’ knowledge may be valuable in designing more
sustainable strategies for controlling pest termites.
Results from recent field surveys in Uganda revealed
significant association between the proportion of farm-
ers practising intercropping in maize and termite
damage severity. During the same surveys, cropping
practices appeared to influence levels of both termite
damage to maize and incidence of predatory ant species.
Based on available literature and on results of the two
surveys, intercropping was considered to be worthy of
detailed evaluation for termite control for subsistence
maize farmers. The objectives of the study were there-
fore to:
(i) determine the effect of intercropping maize with
three legume crops: soyabean, groundnut and
common beans, on severity of termite attack,
(ii) determine the effect of maize–legume intercrops on
the activity of ant species predatory on termites.
2. Materials and methods
This study was conducted at Namulonge Agricultural
and Animal Production Research Institute (NAARI),
during two planting seasons: the second rainy season of
1997 and the first rainy season of 1998. The tests were
conducted in a field with a history of high termite
populations. The experiments were planted on 18
September 1997 and on 7 April 1998, with maize and
the legumes being planted simultaneously.
2.1. Effect of intercropping maize with legumes on
termite damage to maize
Three maize–legume intercrops made up of ground-
nut (var. Red Beauty), soyabean (var. Nam 1) and
Phaseolus beans (var. K31) were established, with sole
maize as the control. The maize variety used, Longe-1, is
known to be susceptible to termite attack (Anon., 1996).
The experimental plots measured 15 m  20 m and
were separated by 1.5 m alleys. Maize was planted at a
spacing of 75 cm between rows and 50 cm between
plants, with 2 plants per hill. In each intercrop plot, two
rows of the respective legume crops were planted
between two rows of maize. Soyabeans were spaced at
25 cm between rows and 12.5 cm between plants while
common beans and groundnuts were spaced at 25 cm
between rows and 25 cm between plants, with one seed
per hole. There were five replicate plots for each
treatment, arranged in a randomised complete block
design.
In one half of each treatment plot lindane (100 g l1
EC), a persistent soil insecticide, was applied 1–2 days
after planting at a rate of 1.5 l ha1 (manufacturers’
recommendation) using a hand-held knapsack sprayer.
The experiment was thus a split-plot design, with
insecticide level as the main-plot factor while the
intercrop type represented the sub-plot factor.
2.2. Sampling for termite damage and predatory ants
Both destructive and non-destructive sampling meth-
ods were used to assess termite damage to maize plants
and nesting of predatory ants. For non-destructive
sampling, 40 plants randomly selected from within the
middle 10 rows of each plot were tagged with blue
ribbon for assessment of termite damage throughout the
season. Each tagged plant was inspected for damage to
stems, leaves and cobs at 14, 28, 42, 56, 70, 84 and 102
days after emergence. Results were expressed as the
proportion of plants showing termite damage out of the
40 sample plants, averaged across the seven sampling
occasions. Termites of the genus Microtermes are known
to damage the root systems of crops. On four occasions,
ten plants were destructively sampled from each sub-
plot to determine if intercropping maize with legumes
affected the activity of root feeding termites. The plants
were uprooted using a hand hoe, caution being taken
not to destroy the root systems. Presence or absence of
termites attacking roots in the different treatments were
recorded four times per season, at seedling, silking,
green cob and dry cob stages, approximately 28, 56, 90
and 105 days after maize emergence.
To assess the incidence of predatory ants in the plots,
the 40 plants sampled for termite damage were inspected
on the same occasions for presence of ant nests within a
25 cm radius around the plant stem bases. An ant-
nesting index expressed as the number of maize plants
out of 40 with ants nests was calculated.
At crop maturity, maize yields were determined by
harvesting cobs from the middle 10 rows of every plot
and extrapolating plot yields to kg ha1.
2.3. Data analysis
Data were tested for conformity to assumptions of
ANOVA as dictated by tests of normality and of
homogeneity of variance (GENSTAT 5 release 4.1).
Data for % plants damaged and ant nesting were pooled
to generate seasonal means and treatment effects
analysed following procedures for factorial designs.
Data on root infestation by Microtermes spp. were
analysed for individual sampling dates. Thereafter, pre-
planned orthogonal comparisons were tested for effects
 B.M. Sekamatte et al. / Crop Protection 22 (2003) 87–93 89

of intercropping and insecticide application on termite
attack, activity of predatory ants, and grain yield. The
means were separated by Duncan’s New Multiple Range
(DMR) test.
3. Results
3.1. Effect of intercropping on termite damage
Analysis of variance for seasonal means of % plants
damaged by termites during the two cropping seasons
showed significant differences among treatments
(Fig. 1). The mean number of damaged plants in the
1997 cropping season varied between 6.2% and 68%
across the treatments (Fig. 1a). Damage was signifi-
cantly (Po0:05) lower in protected plots than in the
unprotected plots in both seasons (Fig. 1). No signifi-
cant differences in % damage were obtained among the
protected crop mixtures.
Termite damage on maize varied with type of legume
intercropped with maize. In the second season of 1997,
80
a b ab
70
% plants damaged
the proportion of damaged plants was very significantly
(Po0:001) greater in sole maize than in maize inter-
cropped with soyabean, but the difference between
damage in sole maize and maize/groundnut or maize/
bean intercrops was not significant (Fig. 1a). In the first
season of 1998, termite damage in sole maize was very
significantly (Po0:001) greater than that in either
maize/soyabean or maize/groundnut intercrops but the
difference between sole damage in maize and in maize/
bean intercrops was not significant (Fig. 1b).
In the second growing season of 1997, there were
o10% of roots infested by Microtermes spp. in all
treatments until 90 days after emergence (DAE). At 105
DAE, there was a significantly (Po0:05) higher
proportion of plant root systems infested by Micro-
termes spp. in sole maize than in either the maize/
groundnut or maize/soyabean intercrops (Fig. 2A). The
pattern of Microtermes infestation of root systems in the
different treatments at 105 DAE, during the first
growing season of 1998, was similar to that in the
previous season but the proportions of infested plants
were approximately half those in 1997 (Fig. 2B). In this
season, the proportion Microtermes infested root
systems was significantly (Po0:05) higher in the sole
maize and the maize/bean treatments than in the
remaining intercrops (Fig. 2B).
ab

60
50 30 a c bc ab
40
% of plants with Microtermes

25
30
20 20
10 15
0
Sole maize Maize/soya Maize/groundnuta Mize/bean 10
(A) Crop combinations protected
unprotected 5

0
60 b Sole maize Maize/soybean Maize/groundnut Maize/beans
a b ab
(A) Crop combinations
% plants damaged

50
40
30 a b b a
30
% plants with Microtermes

25
20
20
10
15
0
Sole maize Maize/soya Maize/groundnut Maize/bean 10
(B) Crop combinations protected 5
unprotected
0
Fig. 1. Proportion of maize plants damaged by termites in insecticide Sole maize Maize/soybean Maize/groundnut Maize/beans
protected and unprotected sole maize and maize intercropped with Crop combinations
(B)
soyabean, groundnut and common bean, respectively, in (A) the
second growing season of 1997 and (B) the first growing season of 1998 Fig. 2. Proportion of plants with roots infested with Microtermes spp.
at Namulonge Research Station, Uganda. Figures are % damage in unprotected sole maize and maize intercropped with soyabean,
averaged over seven sampling occasions across each growing season. groundnut and common bean at 105 days after emergence in (A) the
Columns (unprotected maize) labelled with the same letter do not second growing season of 1997 and (B) the first growing season of
differ significantly. 1998. Columns labelled with the same letter do not differ significantly.
90 B.M. Sekamatte et al. / Crop Protection 22 (2003) 87–93

3.2. Effects of intercrops on nesting of predatory ants
Ant nesting was invariably influenced by the applica-
tion of lindane in all crop combinations (Fig. 3). In the
1997 cropping season, mean nesting index in the lindane
treated plots was generally low and ranged from 0.01 to
0.07 compared to 0.05 to 0.56 in the unprotected plots
(Fig. 3A). The ant-nesting index was also slightly higher
in this cropping season than in the 1998 season (Fig. 3).
During the 1997 season, there were no significant
differences in ant-nesting index between the three
unprotected intercrops, but the index was significantly
(Po0:01) lower in the unprotected monocrop maize
than in any of the intercrops (Fig. 3A).
Ant-nesting indices in unprotected plots during the
1998 cropping season ranged from 0.03 in monocrop
maize to 0.37 in maize/soyabean intercrop (Fig. 3B). As
in the 1997 season, the ant-nesting index was signifi-
cantly (Po0:01) lower in monocrop maize than in
intercropped maize. In this season, the ant-nesting
indices in unprotected maize/soyabean and maize/
groundnut intercrops were significantly (Po0:05)
greater than that in the unprotected maize/bean inter-
crop (Fig. 3B).
There was a significant negative relationship between
% plants damaged and ant nesting in both the first
cropping season (Y ¼ 80:04X þ 56:75 (r2 ¼ 0:731;
P ¼ 0:05) and in the second cropping season
(Y ¼ 32:602X þ 49:2; r2 ¼ 0:57; P ¼ 0:05). Significant
negative relationships were also obtained between ant-
nesting index and the number of plants with cut stems
(r2 ¼ 0:793; df¼ 28; Po0:01) and root damage scores
(r2 ¼ 0:671; df¼ 28; P ¼ 0:05). No significant relation-
ship was obtained between ant nesting and maize plant
lodging.
3.3. Effect on maize yields
There were significant differences (P ¼ 0:05) in maize
grain yields between the intercrops and sole maize in
both seasons. There were also significant differences
between protected and unprotected crops (Fig. 4A). In
the second cropping season of 1997, maize yield in the

4000 b a a ab
0.6 b a a a
3500
0.5
Ant nesting index

3000
0.4
Yield (kg ha-1)

2500
0.3 2000
0.2 1500

0.1 1000

0 500
Sole maize Maize/soya Maize/groundnut Maize/bean
0
(A) Crop combinations protected Sole maize Maize/soybean Maize/groundnut Maize/beans
unprotected (A) Crop combinations
Protected
Unprotected
0.45 c a a b
0.4 3500 b a b b
Ant nesting index

0.35 3000
0.3
Yield (kg ha-1)

2500
0.25
0.2 2000
0.15 1500
0.1
1000
0.05
0 500
Sole maize Maize/soya Maize/groundnut Maize/bean
0
(B) Crop combinations Sole maize Maize/soybean Maize/groundnut Maize/beans
protected
unprotected (B) Crop combinations Protected
Unprotected
Fig. 3. An index of ant nesting in insecticide protected and unpro-
tected sole maize and maize intercropped with soyabean, groundnut Fig. 4. Maize yields (kg ha1) in insecticide protected and unprotected
and common bean, respectively, in (A) the second growing season of sole maize and maize intercropped with soyabean, groundnut and
1997 and (B) the first growing season of 1998 at Namulonge Research common bean, respectively, in (A) the second growing season of 1997
Station, Uganda. See materials and methods for calculation of ant and (B) the first growing season of 1998 at Namulonge Research
index. Columns (unprotected maize) labelled with the same letter do Station, Uganda. Columns (unprotected maize) labelled with the same
not differ significantly. letter do not differ significantly.
 B.M. Sekamatte et al. / Crop Protection 22 (2003) 87–93
unprotected plots of monocrop maize was 24% lower
than in protected crop, whilst mean yield reduction over
the three unprotected intercrops was 15.7%. The
respective yield reductions were 14% for maize/soya-
bean, 15% for maize/groundnut and 18% for maize/
beans. Yield was significantly greater (Po0:05) in
unprotected maize/soyabean and maize/groundnut than
in unprotected sole maize, but yield of maize/bean
intercrops were not significantly different to those in sole
maize (Fig. 4A). There was a yield gain over sole maize
of 12% in the soyabean intercrop, equivalent to
344 kg ha1, and 6% in the maize/groundnut crop,
equivalent to 160 kg ha1.
In the first cropping season of 1998, maize grain yield
in the unprotected maize monocrop was reduced by
23% while yield reduction in the three intercrops
averaged 8%. The respective yield reductions were 8%
for maize/soyabean, 6% for maize/groundnut and 11%
for maize/beans (Fig. 4B). Yield in the unprotected
maize/soyabean intercrop was significantly greater
(Po0:05) than in the sole maize crop but yield in the
other unprotected intercrops did not differ significantly
from that in the sole maize (Fig. 4B). There was a yield
gain over sole maize of 19% in the maize/soyabean
intercrop, equivalent to 418 kg ha1.
4. Discussion
These results have demonstrated that intercropping
maize with a range of food legumes is associated with a
significant reduction in termite attack on maize and an
increase in the nesting of predatory ants in maize fields
under the conditions prevailing at the study site. The
impact of the practice depended largely on the type of
legume intercropped with maize. Termite attack was
lower in the maize/soyabean intercrop than in maize
intercrops with groundnut and beans. This observation
was probably related to maturity time and to the
amount of foliage litter shed over the growth period of
the different legume species. The bean variety used in the
study (K131) attained physiological maturity in 70–75
days while the groundnut and soyabean varieties
matured in about 120 days, approximately the same
maturity period as for Longe-1 maize. Species of the
genera Macrotermes and Pseudacanthotermes, which
were particularly important termite pests in experimen-
tal plots, were observed to feed actively on the leaf litter
shed by the legumes, particularly for groundnut and
soyabean. Both these lasted longer than the beans in the
field and provided more food to termites. The soyabean
variety (NAM-1) shed considerable amounts of foliage,
especially from pod filling through pod maturation
phases, and the additional litter available to termites
was presumed to partially account for the reduced
damage to maize in this crop combination. This
91
preference of dry foliage by Macrotermitinae was also
reported in previous studies by Bigger (1966) and Sands
(1977).
Similar effects on levels of termite attack were
observed in related studies on the effects of mulching
maize with different quantities of stover. Termite attack
on maize was negatively correlated with the quantity of
mulch applied in each treatment (Sekamatte et al.,
2001). The reduction in termite attack on maize in the
intercrops may also have been affected by the thick
legume canopy in the intercrops. Although there is no
reported crop-based study on this phenomenon
relating to termites, similar observations were made in
a forest system (Tho, 1974) indicating that attack by
Macrotermes sp. on Eucalyptus ceased after canopy
closure and also that populations of these termites
disappeared under mature monocultures of Eucalyptus
plantations.
We recorded significantly higher numbers of nests of
predatory ants in the intercrops compared to monocrop
maize. Similar high levels of ant nesting were observed
when maize was mulched with stover (Sekamatte et al.,
2001). Increased abundance of ground-active natural
enemies have been reported in maize-groundnut mix-
tures in the Philippines (Perfecto and Sediles, 1992) and
in cereals in the UK (Potts and Vickerman, 1974),
presumably associated with increased humidity and
shade near the soil. This modification of microclimate
could account for the increased ant nesting in intercrops
in the present study. While it is likely that both the
improvements in termite food sources in the form of dry
legume foliage and enhanced predation by ants
accounted for the observed reduction in termite attack
on maize plants, it was not possible to distinguish the
effects of the two mechanisms.
Although Myrmicaria spp. have been widely recog-
nised as important ground predators (Perfecto, 1990;
Kenne and Dejean, 1999; Kenne et al., 2000), we have
found no published work on the predatory role of
Lepisiota species, despite the fact that they are widely
known and valued by farmers in Uganda, especially in
the cotton-based cropping systems in northern Uganda.
Lepisiota ants were actively seen carrying termites larger
than themselves in the experimental plots. The ants also
nest extensively around the bases of maize plants,
presumably making it difficult for termites to attack
such protected plants. However, the ecological role
(Risch and Carrol, 1982) and the potential of ground-
active ants in the control of a range of crop pests has
been reported elsewhere (Lim, 1990; Perfecto and
Sediles, 1992).
The yield advantage of intercropping maize with
soyabean was relatively small, 12% in the second season
of 1997 and 19% in the first season of 1998. However,
these figures, equivalent to 344 and 418 kg ha1,
respectively, could be significant to smallholder farmers
92 B.M. Sekamatte et al. / Crop Protection 22 (2003) 87–93
in Uganda for whom yields rarely exceed 1.5 t ha1. A
yield advantage (6%) from intercropping maize with
groundnut was only recorded in the second growing
season of 1997. In principle, improved maize yields
in the intercrops might have been partially due to
factors other than reduced termite damage, such as
better maize growth resulting from improved nitrogen
status of soils in plots with legumes. However, the fact
that yields in the protected intercrops were either the
same as or in some cases lower than, those in the sole
maize crop suggests that this was not an important
factor.
Sands (1977) and Tho (1974) viewed intercropping as
a means of preventing specific termite species, achieving
pest status in forestry through maintaining species
diversity. Lee and Wood (1971) reported disappearance
of Nasutitermes exitiosus (Hill) when Pinus sp. were
mixed with Eucalyptus species in Australia. Cases have
also been reported where establishment of monoculture
forestry plantations has led to the build-up of some
termite species (e.g. Coptotermes sp.) in southeast Asia
(Logan et al., 1990). Certain grasses are grown in West
Africa to repel termites (Malaka, 1972), although the
effects of intercrops on termite damage in field crops are
still, largely, poorly understood. Gold and Whightman
(1991) and Hillocks et al. (1996) considered intercrop-
ping and mulching among potentially useful cultural
control options for subsistence farmers, but pointed out
that they were still largely untested.
In conclusion, maize–legume intercrops may, in
appropriate circumstances, help to suppress termite
damage and to increase grain yield of maize. Different
species of legumes appear to differ in their influence on
termite damage and crop loss. In the present study, two
legumes, soyabean and groundnut, suppressed termite
attack and improved yield of maize better than common
beans. The greatest impact of intercropping is likely to
be in areas where termite infestation is currently a major
production constraint. It is possible that other legumes,
or indeed other crops (e.g. sorghum), could be used for
this purpose. In areas where smallholder farming
systems include livestock, forage legumes might be
included as an intercrop. The willingness of smallholder
farmers to adopt new inter-cropping practices will be
determined as much by socio-economic factors as by
purely technical constraints.
Acknowledgements
We thank our colleagues James Akono, James
Kayongo and Solomon Kaboyo for their dedicated
participation at all stages of these experiments. This
paper is an output from a research project (R6653)
funded by the United Kingdom, Department for
International Development (DFID).
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