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Copyright © 2009 by the author(s). Published here under license by the Resilience Alliance.
Sileshi, G. W., P. Nyeko, P. O. Y. Nkunika, B. M. Sekematte, F. K. Akinnifesi, and O. C. Ajayi. 2009.
Integrating ethno-ecological and scientific knowledge of termites for sustainable termite management and
human welfare in Africa. Ecology and Society 14(1): 48. [online] URL: http://www.ecologyandsociety.org/
vol14/iss1/art48/
Insight
Integrating Ethno-Ecological and Scientific Knowledge of Termites for
Sustainable Termite Management and Human Welfare in Africa
Gudeta W. Sileshi
1
, Philip Nyeko
2
, Phillip O. Y. Nkunika
3
, Benjamin M. Sekematte
4
,
Festus K. Akinnifesi
1
, and Oluyede C. Ajayi
1
ABSTRACT. Despite their well-known role as pests, termites also provide essential ecosystem services.
In this paper, we undertook a comprehensive review of studies on human–termite interactions and farmers’
indigenous knowledge across Sub-Saharan Africa in an effort to build coherent principles for termite
management. The review revealed that local communities have comprehensive indigenous knowledge of
termite ecology and taxonomy, and apply various indigenous control practices. Many communities also
have elaborate knowledge of the nutritional and medicinal value of termites and mushrooms associated
with termite nests. Children and women also widely consume termite mound soil for nutritional or other
benefits encouraged by indigenous belief systems. In addition, subsistence farmers use termites as indicators
of soil fertility, and use termite mound soil in low-risk farming strategies for crop production. In the past,
chemical control of termites has been initiated without empirical data on the termite species, their damage
threshold, and the social, ecological, or economic risks and trade-offs of the control. This review has
provided new insights into the intimate nature of human–termite interactions in Africa and the risks of
chemical control of termites to human welfare and the environment. We recommend that management of
termites in future should be built on farmers’ indigenous knowledge and adequate understanding of the
ecology of the local termite species.
Key Words: Agroforestry; biodiversity; geophagy; management; traditional ecological knowledge
INTRODUCTION
A distinct dichotomy exists between the pest
management literature that depicts termites as
“pests” and the ecological literature demonstrating
their crucial role in ecosystems. There is no doubt
that some species cause significant damage to crops,
rangeland, trees, and structural timber. At the same
time, they also play a beneficial role through
promotion of essential ecological processes. The
ongoing interest in sustainable agriculture and food
security in Africa highlights the need for a more
balanced approach to termite control and
maintenance of their ecosystem services. To begin
to address the mismatch between these two
objectives, a holistic appraisal of the termite
problem and opportunities for their sustainable
management is needed. Sustainable termite
management is defined here as one that ensures (1)
control of the pest species without causing
ecological damage and loss of the ecosystem
services provided by termites, (2) conservation of
the non-pest termite species, and (3) use of termites
and associated resources without exhausting them.
Management of risk and ensuring resilience are key
concepts in sustainability, and these beg for a
strategy that combines the skills and indigenous
technical knowledge of farmers with modern
scientific knowledge (Sileshi et al. 2008a).
Termites are a large and diverse group of insects
consisting of over 2600 species worldwide. With
over 660 species, Africa is by far the richest
continent in termite diversity (Eggleton 2000). Most
(ca. 85%) of the known genera and those species
that damage crops, trees, and rangeland belong to
the family Termitidae. This family consists of four
subfamilies: Macrotermitinae, Nasutitermitinae,
Termitinae, and Apictotermitinae. Although the
exact number of pest species is not known, it is
estimated that less than 20% of members of the
family Termitidae are serious pests (Pomeroy et al.
1
World Agroforestry Centre (ICRAF),
2
Makerere University,
3
University of Zambia,
4
Nikoola Institutional Development Associates
Ecology and Society 14(1): 48
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1991, Mitchell 2002). Over 90% of the termite
damage in agriculture, forestry, and urban settings
is attributed to members of the Macrotermitinae
(Pomeroy et al. 1991, Mitchell 2002), which build
the large mounds (hereafter called termitaria) that
form the spectacular features of the African
landscape (Glover 1967, Malaisse 1978). The
reputation of termites as pests is also associated with
the presence of termitaria in crop fields and near
trees. Thus, most of the discussion in this paper
focuses on the members of the Macrotermitinae.
The main genera in the subfamily Macrotermitinae
are Odontotermes (79 species), Microtermes (33
species), Macrotermes (21 species), Ancistrotermes
(nine species), Allodontermes (seven species), and
Pseudacanthotermes (five species) (Pomeroy et al.
1991, Darlington et al. 2008). The taxonomy of
Macrotermitinae is notoriously difficult, and many
species are not easy to identify with certainty
(Darlington et al. 2008). Unfortunately, current
recommendations for termite control do not take
into account this taxonomic difficulty.
Macrotermes, Odontotermes, and Microtermes
species tolerate semi-arid and even arid areas
(Eggleton 2000). Forty percent of Africa’s
population lives in arid, semi-arid, and dry sub-
humid areas (see map by D. Digout on the United
Nations Environment Programme (UNEP) GRID-
Arendal website
http://maps.grida.no/go/graphic/ar
idity_zones
), and practices agro-pastoral production
systems. In traditional agro-pastoral systems, when
land productivity declines, the farmers shift to a new
area. With the rapid increase in human population,
the rate of conversion of natural habitat has
increased and land degradation is taking place at a
faster rate (United Nations University (UNU)
1998). Approximately 22% of vegetated land in
Africa has been classified as degraded, and 66% of
this is classified as moderately to severely degraded
(UNU 1998). Most of these areas experience
periodic droughts, and climate change has made
them even more prone to frequent dry spells. Crops
and trees are increasingly planted on marginal land,
resulting in greater stress and vulnerability to
termite attack (Glover 1967, Wood et al. 1980). In
some places, such as the “Cattle Corridor” of
Uganda, overstocking has led to depletion of forage
to the extent that termites have to feed on whatever
lies in their way (Sekamatte and Okwakol 2007,
Tenywa 2008). As humans encroach into bushland,
the conflict between humans and pest termites will
increase.
It must be noted that the pest activity is a part and
parcel of the termite’s beneficial role in various
ecosystems (Glover 1967, Black and Okwakol
1997). Macrotermitinae collect up to 60% of the
grass, woody material, and annual leaf fall to
construct the fungus gardens in their nests (Lepage
et al. 1993). This translates to 1.5 tons of litter
removal per hectare every year (Jones 1990, Lepage
et al. 1993), which leads to a dramatic reduction in
“fuel load” and fire intensity, while at the same time
preserving nutrients in termitaria beyond the reach
of fire (Lepage et al. 1993). By breaking down up
to 100% of the litter fall and mineralization of up to
50% of the net primary production, termites
influence ecosystem services such as nutrient
cycling and biomass production (Jones 1990). From
this viewpoint, termites are beneficial for the
functioning of forest and savannah ecosystems.
Termites also play a significant role in the
availability of nutrients and water to crops and trees,
and hence the productivity of agricultural
ecosystems (Black and Okwakol 1997). Throughout
the semi-arid regions of West Africa, crop-growth
variability related to termite activity has been used
by subsistence farmers in low-risk farming
strategies for crop production (Mielke and Mielke
1982, Brouwer et al. 1993, Harris et al. 1994).
Therefore, agriculture holds the key to the
management of termites and conservation of
biodiversity as small-scale farmers are the ultimate
managers and stewards of the land (Sileshi et al.
2008a). Resource-poor farmers look for practices
that best fit their biophysical, economic, and
sociocultural conditions. Ethno-ecology or traditional
ecological knowledge (Berkes 2008) is particularly
important for the formulation of sustainable termite
management in Africa. Traditional ecological
knowledge is defined as “a cumulative body of
knowledge, practice and belief evolving by adaptive
processes and handed down through generations by
cultural transmission, about the relationship of
living beings with one another and with their
environment” (Berkes 2008). Engagement of
researchers with local communities may thus help
researchers to link their efforts to the local
environmental and cultural context.
The objective of this paper is to bring together
farmers’ ethno-ecological knowledge and evidence
from soil science and ecology in order to build more
coherent principles for sustainable termite
management. Toward that end, we undertook a
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comprehensive review of case studies on human–
termite interactions from across Sub-Saharan Africa
(Fig. 1) and, summarize the key findings in a
comparative framework (indigenous vs. scientific)
using the following specific themes: (1) farmers’
ethno-ecological knowledge of termites and (2)
termite management practices. Finally, we describe
the potential impacts of termite management
practices on the ecosystem services that termite
species provide in the agro-pastoral systems of Sub-
Saharan Africa.
METHODS
More than 10 studies related to farmers’ ethno-
ecological knowledge of termites were found by
searching the internet and databases, and checking
the references of published studies across Sub-
Saharan Africa. Emphasis was placed on eastern
and southern Africa because this is the geographic
area where the Macrotermitinae, which are the most
serious pests in agriculture and forestry, reach their
highest densities (Jones 1990). The bulk of the
discussion is based on studies conducted in five
districts of Malawi (Sileshi et al. 2008b), two
districts of northwest Mozambique (Sileshi et al.
2008b), three districts of eastern Zambia (Sileshi et
al. 2008b), six districts of southern Zambia
(Nkunika 1998), Tororo district of Uganda (Nyeko
and Olubayo 2005), and Machakos district of Kenya
(Malaret and Ngoru, 1989). The specific sites (Fig.
1) are characterized by extreme variability of
rainfall, drought, and dry spells. Drought is defined
as absence of rainfall over an extended period of
time, usually a season or more, whereas dry spell is
a period of abnormally dry weather conditions less
severe than those of a drought.
The data-collection methods employed in the
specific study sites (Fig. 1) included community
meetings, focus-group interviews, application of
semi-structured questionnaires to farmers, and
direct field observation by the authors. A
participatory rural appraisal (PRA) was used in the
study in eastern Uganda (Nyeko and Olubayo 2005).
The farmers in Uganda gave information on (1)
identity of the termite species, (2) abundance and
distribution of termites, (3) termite damage to trees
and crops, (4) relationship between termite damage
and rainfall patterns, (5) priority termite species for
control, (6) termites as a source of food, and (7)
termite control practices. During community
meetings and focus-group interviews in Malawi,
Mozambique, and Zambia (Nkunika 1998, Sileshi
et al. 2008b), farmers provided information on (1)
cultivated plants attacked by termites, (2) stage of
crop susceptible to attack, (3) time of termite attack,
and (4) indigenous control practices. During group
interviews in Kenya, farmers provided information
on (1) local names and descriptions of termites, (2)
soil and land-use preferences of termites, (3)
cultivated and wild plants attacked by termites, (4)
stage and condition of the plants when attacked, (5)
termite-resistant or repellent plants, and (6) termite
control practices (Malaret and Ngoru 1989).
Individual interviews designed to supplement
results from community meetings were also
conducted in eastern Zambia. In all study sites, the
researchers made direct field observations,
collected specimens to confirm termite identity or
their damage, and obtained authoritative identification
of the termite samples.
SYNTHESIS
Farmers’ Ethno–Ecological Knowledge
Review of the specific studies and the general
literature indicated that farmers have a rich
knowledge of termite biology and the ecosystem
services provided by termites. What follows is a
brief summary of key findings using a comparative
analysis of farmers’ indigenous knowledge and
scientific knowledge.
Farmers’ knowledge of termite taxonomy
Local communities are able to identify the major
genera and species using vernacular names (Table
1). For example, in Tororo district of Uganda, a total
of 14 species were identified with distinct
vernacular names (Nyeko and Olubayo 2005; Table
1). Similarly, 70%–100% of the farmers
interviewed across five other districts of Uganda
were able to identify various termites in their area
(Sekamatte and Okwakol 2007). Farmers were also
able to identify termites using vernacular names in
Kenya (Malaret and Ngoru 1989; Table 1), Somalia
(Glover 1967), Zambia, and Malawi (Nkunika
1998, Sileshi et al. 2008b; Tables 1, 2). Farmers’
taxonomy also showed high internal consistency
and closely corresponded to termite genera (Malaret
and Ngoru 1989, Nyeko and Olubayo 2005).
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Fig. 1. The specific study sites and countries in Sub-Saharan Africa (in black outline) for which
literature (L) was available for this synthesis. The specific sites are (1) Tororo district in eastern Uganda,
(2) Machakos district in eastern Kenya, (3) Kasungu, (4) Mchinji, (4) Dedza, (5) Ntcheu, (6) Balaka, and
(7) Zomba districts in Malawi, (8) Angonia and (9) Tsangano districts in northwest Mozambique, (10)
Chipata and (11) Katete in eastern Zambia, (12) Mazabuka and Monze, (13) Gwembe and Choma, and
(14) Kalomo and Livingstone districts in southern Zambia. The study sites were superimposed on the
aridity map of Africa (UNEP/GRID-Arendal 2002).
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Table 1. List of termite genera and species, their corresponding vernacular names, indigenous knowledge
of their pest status, and uses as sources of food and mushrooms in Zambia, Malawi, Uganda, and Kenya
Country District Species name (local name) Vernacular name Pest Edible Mushroom
Zambia
a
Kalomo Macrotermes falciger Machenya Yes Yes Yes
Gwembe Muchenje, Chaca, Machenya Yes Yes Yes
Mazabuka Muchenje, Makwakwasi Yes Yes Yes
Monze Muchenje, Chisinza Yes Yes Yes
Choma Simachenya, Machenya Yes Yes Yes
Livingstone Machekenene, Mpango Yes Yes Yes
Monze Odontotermes spp. Chuulu, Mumona Yes No Yes
All Allodontotermes spp. Lumona, Mulyazi, Chimuma Yes No No
Microtermes spp. Lumona, Mulyazi, Chimuma Yes No No
Amitermes truncatidens Chuulu, Mumona Yes No No
Pseudacantthotermes sp. Lumona, Mulyazi, Mulahi Yes Yes Yes
Trinervitermes rhodesiensis Lumona, Mulyazi No No No
Monze Cubitermes tenuiceps Kashimbwa, Tuumbusu No No No
Zambia
b
Chipata Macrotermes subhyalinus Kalanzi Yes Yes Yes
Chipata M. falciger Magenge Yes Yes Yes
Chipata Odontotermes spp. Gegedule Yes Yes ND
Microtermes spp. Kauni Yes ND ND
Malawi
b
Balaka M. michaelseni Madulila Yes Yes Yes
Uganda
c
Tororo Macrotermes subhyalinus Agoro Yes Yes Yes
Macrotermes bellicosus Ripo Yes Yes No
Cubitermes ugandensis Aming No No No
Basidentitermes sp. Aming No No No
Microcerotermes sp. Kithea No No No
Odontotermes kibarensis Magrere Yes Yes Yes
Odontotermes (?latricius Singiri No Yes Yes
Odontotermes sp. 1 Ogwee Yes Yes Yes
(con'd)
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Odontotermes sp. 2 Mbala No Yes No
Pseudacanthotermes spriniger Miyal Yes Yes Yes
Pseudacanthotermes militaris Sisi Yes Yes Yes
Pseudacanthotermes sp. Wambwe Yes Yes Yes
Amitermes (? Truncatidens Rudho Yes No No
Trinervitermes oeconomus Thuk No No No
Kenya
d
Machakos Macrotermes spp. Nthwa, Mungasa Yes ND ND
Odontotermes spp. Mbaa, Mbaawa Yes ND ND
Microcerotermes spp. Mungumi, Kathoa Yes ND ND
Pseudacntotermes spp. Ngai, Kikai Yes ND ND
Allodontotermes spp. Ndulamatu, Mungasa Yes ND ND
Cubitermes spp. Kikii No ND ND
a
Nkunika (1998);
b
Sileshi et al. (2008b);
c
Nyeko and Olubayo (2005);
d
Malaret and Ngoru (1989)
? Species name uncertain
No = termite does not produce mushrooms or the mushroom produced is not edible
ND = not determined
Farmers’ knowledge of termite abundance and
distribution
Farmers have also good understanding of the
abundance and distribution of termites in their area.
For example, most farmers in Tororo district rated
Macrotermes bellicosus and M. subhyallinus as the
most abundant (Nyeko and Olubayo 2005). This is
in agreement with scientific studies in Uganda
(Pomeroy 1978). However, farmers’ rating of M.
bellicosus as more abundant in the upland and M.
subhyallinus more abundant in the lower altitudes
disagrees with Pomeroy (1978), who states that the
two species have similar distributions. Farmers in
Machakos district of Kenya associated Macrotermes
and Odontotermes with farmland more than
bushland. They could also identify the humus
feeders from those that attack crops or trees (Malaret
and Ngoru 1989).
Farmers’ perception of termites as pests
Studies evaluating farmers’ perceptions of termites
(Nkunika 1998, Nyeko and Olubayo 2005,
Sekamatte and Okwakol 2007, Sileshi et al. 2008b)
have demonstrated that farmers have good
knowledge of those species that are pests (Tables 1,
2). In Tororo district of Uganda, farmers identified
eight pest species (Table 1). However, they rated M.
bellicosus and M. subhyallinus as more serious than
the other species (Nyeko and Olubayo 2005).
Although Pseudacanthotermes spriniger was
reported to damage some crops, farmers perceived
it as a minor pest that does not merit control (Nyeko
and Olubayo 2005). Out of the six genera identified
by farmers in southern Zambia, M. falciger was at
the top of the pest list (Nkunika 1998). In eastern
Zambia, farmers ascribed most of the crop damage
to M. falciger and M. subhyallinus (Table 2).
Microtermes species considered major pests in
Africa (Wood et al. 1980) were not rated as serious
pests by most farmers (Malaret and Ngoru 1989,
Nyeko and Olubayo 2005, Sileshi et al. 2008b). This
is probably because Microtermes species do not
build termitaria. Damage to plants by Microtermes,
Allodontermes, and Ancistrotermes spp. is often
internal or subterranean. This probably makes these
species less apparent to farmers.
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Table 2. Percentage of respondents identifying termites that attack crop plants by local names in eastern
Zambia (source: Sileshi, unpublished data)
Number of
respondents
a
Percentage of cases where the termite’s local name was mentioned or unknown
Magenge Kalanzi Muswe Chiswe Gegedule Mphusi Kauni Unknown
Crop
Maize 87 (100) 45.3 30.5 11.6 5.2 1.1 0 1.1 5.3
Groundnut 78 (89.7) 37.8 34.6 12.7 5.1 1.2 1.3 1.3 6.1
Sunflower 24 (27.6) 45.8 33.3 12.5 8.3 0 0 0 0
Cotton 23 (26.4) 47.8 17.4 13.0 8.7 0 4.4 0 8.7
Soybean 19 (21.8) 68.4 21.1 5.3 0 0 0 0 5.3
Beans 17 (19.5) 41.2 35.3 11.8 0 5.9 0 0 5.9
Cassava 12 (13.8) 58.3 25.0 0 8.3 0 0 0 8.3
Cowpea 3 (3.4) 33.4 66.6 0 0 0 0 0 0.0
a
Figures in parentheses represent percentage of respondents out of a total of 87
Farmers’ perception of susceptibility of plants to
termites
Farmers were also able to identify which crop or
tree species were susceptible to termite attack and
which were resistant. Farmers rated maize (Zea
mays L.), groundnut (Arachis hypogaea L.), and
sugarcane (Saccharum spp.) as highly susceptible
to termite damage in Uganda (Nyeko and Olubayo
2005), Kenya (Malaret and Ngoru 1989), and
Zambia (Sileshi et al. 2008b). Maize and groundnut
were particularly rated as highly susceptible by most
respondents (>90%) in eastern Zambia (Table 3).
The farmers’ assessment is in agreement with the
entomological studies conducted elsewhere in
Africa (e.g., Wood et al. 1980). In Machakos district
of Kenya, farmers identified 24 species of trees and
shrubs as resistant to termites (Malaret and Ngoru
1989). Ugandan farmers identified Eucalyptus
species and Grevillea robusta as particularly highly
susceptible to M. bellicosus and M. subhyallinus
(Nyeko and Olubayo 2005). The more serious
damage on the exotic crops (e.g., maize and
groundnuts) and trees (e.g., Eucalyptus and
Grevillea) is probably because these species lack
resistance to African termites (Logan et al. 1990,
Mielke and Mielke 1982). Indigenous African crops
and trees are expected to be resistant to these
termites with which they have co-evolved. Contrary
to scientific reports that termites rarely attack
vigorously growing plants (Wood et al. 1980), some
farmers stated that termites attack both healthy and
unhealthy plants (Malaret and Ngoru 1989). This
does not necessarily mean that farmers are incorrect
in their assessment. Rather it indicates that there are
areas where local knowledge differs from scientific
studies.
Most farmers in eastern Zambia believe that crops
become susceptible to termite attack at maturity
(Table 3), and this agrees with the literature (Wood
et al. 1980). According to these farmers, termite
damage to crops and trees is more severe during dry
spells or drought periods. Similarly, Ugandan and
Kenyan farmers considered termite damage to be
more severe in the dry months compared with the
wet months (Malaret and Ngoru 1989, Nyeko and
Olubayo 2005). Damage by termites is greater
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Table 3. Farmers’ perception of susceptibility of crops to termite damage and time of crop damage in
eastern Zambia (source: Sileshi, unpublished data)
Rating of crop as
susceptible (%)
a
Rating of time of crop damage (% of respondents
b
)
Continuous Dry spell
(rainy season)
Dry
season
Do not
known
Period of highest damage
Crop
Maize 100.0 (87) 2.1 88.4 8.4 1.1 Flowering onward (83.1%)
Groundnut 89.7 (78) 0 84.6 12.8 2.6 Flowering onward (82.1%)
Sunflower 27.6 (24) 0 70.8 29.2 0 Flowering onward (85.0%)
Cotton 26.4 (23) 4.4 95.6 0 0 Flowering onward (69.5%)
Soybean 21.8 (19) 10.5 73.7 15.8 0 Flowering onward (73.7%)
Beans 19.5 (17) 17.7 76.5 0 5.9 Flowering onward (65.7%)
Cassava 13.8 (12) 33.3 33.3 33.3 0 Maturity (41.7%)
Sugar cane 4.6 (4) 25.0 0 75.0 0 Throughout the year (100%)
a
Figures in parentheses represent the total number of respondents
b
The % was calculated using the respective number of respondents for each crop (figures in parentheses
in column 2)
during dry periods or droughts than periods of
regular rainfall (Logan et al. 1990, Black and
Okwakol 1997, Sileshi et al. 2005). The increases
in termite damage could also be associated with
climate change-induced drought. In recent decades,
drought linked to El Niño episodes has become more
intense and widespread in southern Africa
(Fauchereau et al. 2003).
Farmers in Uganda and Zambia also mentioned that
termite problems are more serious now than in the
past (Sekamatte and Okwakol 2007, Sileshi et al.
2008b, Tenywa 2008). In a recent survey of eastern
Uganda, elders linked the increasing termite
problem and low abundance of predatory ant species
to aerial sprays intended to control tsetse flies
(Glossina sp.) during the 1960s and 1970s
(Sekamatte and Okwakol 2007). Termite damage
on trees and crops could have increased as a result
of the depletion of the usual termite food due to
deforestation and overgrazing (Tenywa 2008).
Deforestation may also have resulted in loss of the
natural enemies of termites such as the aardvark
(Orycteropus afer), pangolin (Manis spp.), aardwolf
(Proteles cristatus), and hedgehog tenrec (Echinops
tefairi) (Pomeroy et al. 1991, Peveling et al. 2003).
Habitat loss has led to the disappearance of the
aardvark in countries such as Ethiopia (Jiru 2006).
Continuous cultivation and overstocking reduce the
diversity of termites (Glover 1967, Black and
Okwakol 1997, Eggleton et al. 2002) followed by
outbreak of those species that tolerate disturbances
(Wood et al. 1980).
Farmers’ knowledge of the role of termites in
human nutrition and health
The literature reviewed indicated that farmers have
extensive knowledge of the value of termites in
human nutrition. The alates, queen, and soldiers of
some species in the subfamily Macrotermitinae are
eaten across most of Africa. Local people easily tell
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the edible termites from those unsuitable for
consumption (Table 1). Out of the 14 species
identified in Tororo district, only 10 were
consumed. Some species were not eaten due to
various reasons (Nyeko and Olubayo 2005).
Some communities also have an intimate
knowledge of the association between termites and
edible mushrooms (Oso 1975, Yongabi et al. 2004,
Nyeko and Olubayo 2005, Kabasa et al. 2006). The
Macrotermitinae cultivate edible mushrooms in the
genus Termitomyces in the termitaria. Each year,
these fungi produce a crop of large mushrooms,
which are highly prized by people as a delicacy.
According to the Guinness Book of Records, T.
titanicus, which grows on termitaria, is the world’s
largest and one of the tastiest mushrooms.
Communities in Uganda and Nigeria associated
specific mushrooms with certain termite species and
identified each Termitomyces species by local name
(Oso 1975, Nyeko and Olubayo 2005). Many
communities in Africa have elaborate indigenous
knowledge of the nutritional and medicinal values
of these mushrooms (Oso 1975, Yongabi et al. 2004,
Nyeko and Olubayo 2005, Kabasa et al. 2006).
According to an indigenous knowledge survey in
Uganda, 45% of the respondents consumed T.
microcarpus for its medicinal value, whereas 5%
consumed it just as food. Chemical analyses
(Kabasa et al. 2006, Opige et al. 2006) demonstrate
that Termitomyces species provide a low-fat, high-
fiber diet rich in protein, minerals (Ca, P, K), and
vitamins. In Cameroon, T. titanicus is dried and
mixed with pastry for the consumption of
underweight children (Yongabi et al. 2004).
Termitomyces species are also used in the treatment
of various diseases (Kabasa et al. 2006).
Termites also contribute to human nutrition and
health through the deliberate ingestion of soil, a
phenomenon called geophagy (Hunter 1973).
Geophagy has been practiced by humans since
antiquity and on almost every continent (Hunter
1973, Rowland 2002). Termitaria are the major
sources of the soil consumed by women and children
in Kenya (Geissler 2000, Luoba et al. 2004),
Tanzania (Knudsen 2002), Zambia, Zimbabwe
(Nchito et al. 2004), and South Africa (Saathoff et
al. 2002). In many communities, geophagy is firmly
grounded in indigenous belief systems (Geissler
2000, Knudsen 2002). In the cosmology of the Luo
(a tribe in eastern Kenya) and Chagga (a tribe in
northern Tanzania), termitaria symbolize sexuality
and the feminine power of procreation, thus
ingesting soil is good for the blood and for fertility
of the woman (Geissler 2000, Knudsen 2002).
Similarly, in cosmology of the San people in
southern Africa, termitaria symbolize a spirit world
that has unparalleled transformative and generative
powers (Mguni 2006). Consumption of soil from
termitaria is common among the nutritionally
vulnerable populations, especially children and
pregnant and lactating women (Wiley and Katz
1998). In a study involving 827 women in Bondo
district of Western Kenya, 46% reported geophagy
during pregnancy and lactation (Luoba et al. 2004).
Among children aged 5–18 in Western Kenya and
Zambia, 73%–74% had ingested soil (Geissler
2000, Nchito et al. 2004). Consumption of soil was
reported more often in girls (53%–80%) than in boys
(39%–68%) in Zambia (Nchito et al. 2004) and
South Africa (Saathoff et al. 2002). Among the
positive health benefits of geophagy during
pregnancy are improved maternal calcium status,
improved foetal skeletal formation and birth weight,
reduction in pregnancy-induced hypertension, and
decreased risk of embryonic exposure to teratogens
and loss of nutrients through emesis (Wiley and
Katz 1998). Another adaptive function of
consuming termite soil is the ability of clays to
adsorb toxins from plants eaten by humans. Soil
from termitaria is high in kaolinic clays, the same
base used in Kaopectate, which is prescribed for
stomach upset. This indigenous knowledge,
therefore, has significant implications for
adaptation to many of the toxins in plants eaten by
people (Rowland 2002).
Farmers’ knowledge of the role of termites in soil
fertility
Subsistence farmers largely base their nutrient-
management strategies on their perception of niche
fertility (Brouwer et al. 1993, Adamou et al. 2007,
Chikuvire et al. 2007). Within-field variability of
soil properties creates niches that farmers perceive
as essential to their farming (Brouwer et al. 1993).
Termites, specifically the Macrotermitinae, create
such niches and in most cases help farmers to reduce
risks of crop failure (Mielke and Mielke 1982,
Brouwer et al. 1993, Harris et al. 1994). Thus,
farmers use termites as biological indicators of soil
fertility status. Traditional soil classification is also
based on termite presence or absence among some
communities (Bellon et al. 1999, Ettema 1994,
Adamou et al. 2007). Therefore, termites are an
important element in traditional land-management
practices in the Sahel (Mando et al. 1999), East
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Africa, and southern Africa (Mielke and Mielke
1982, Nyamapfene 1986). Subsistence farmers
across Sub-Saharan Africa also encourage termite
activity in their crop fields. For example, farmers in
Burkina Faso bury manure in holes near newly
planted millet in the belief that termites attracted to
the manure will improve water availability to the
crop (Logan 1992). Quantitative studies in Burkina
Faso have demonstrated that this practice increases
infiltration rates (Mando et al. 1999). Studies
elsewhere also have demonstrated positive impact
of termites on soil properties and water dynamics
(Chikuvire et al. 2007).
Farmers’ use of termite-modified soil in crop
production has been documented in Uganda
(Okwakol and Sekamatte 2007), Zambia (Siame
2005), Zimbabwe (Bellon et al. 1999, Nyamapfene
1986), Niger (Brouwer et al. 1993), and Sierra
Leone (Ettema 1994). Farmers either plant specific
crops on termitaria (Fig. 2) or spread soil from
termitaria in their fields (Nyamapfene 1986, Logan
1992). A typical example of cropping around
termitaria is the chitemene agriculture in
southwestern Tanzania (Mielke and Mielke 1982).
In Malawi, farmers plant various crops including
bananas (Musa spp.) around termite mounds (Fig.
2). Ugandan farmers plant pumpkins (Cucurbita
spp.), tomatoes (Solanum spp.), onions (Allium
spp.), and maize adjacent to termitaria (Okwakol
and Sekamatte 2007). In Zimbabwe, crops such as
okra (Abelmoschus esculentus), pumpkins, sweet
sorghum (Sorghum spp.), and late-season maize,
which require good water and nutrient supply, are
grown almost exclusively on termitaria (Nyamapfene
1986). In Niger, farmers preferentially plant
sorghum on termitaria (Brouwer et al. 1993). In
some areas, farmers break termitaria and spread the
soil in their field. For example, in southern Zambia,
farmers remove portions of the mound, making sure
that they leave the base intact so that the colony is
not destroyed. This soil is then transported to the
field and worked into the top soil before the rains
begin. In areas where conservation farming is
practiced, soil from termite mounds is placed into
planting basins (Siame 2005). In South Africa,
circular patches of exceptionally well-grown
sugarcane, known as “isiduli”, are a common
feature of sandy sugarcane fields. These correspond
to mounds of M. natalensis leveled by ploughing
(Cadet et al. 2004). Similarly, in Zimbabwe, farmers
widely use soil from termitaria to improve soil
fertility (Bellon et al. 1999, Nyamapfene 1986).
These farmers’ practices have been a subject of
studies, and scientific explanations do exist for most
of them (Watson 1977, Nyamapfene 1986). For
example, detailed studies show that cane yield is
five times greater using the “isiduli” than elsewhere
in the field (Cadet et al. 2004). Similarly, plant
biomass and grass growth were significantly higher
around termitaria compared with the open veld in
the Eastern Cape (Steinke and Nell 1989). The
increased growth of grass surrounding termitaria
was due largely to accumulation of runoff water at
the base. Not only could this lead to increased
productivity during dry years, but it could also make
it possible for plants to survive intense drought
(Steinke and Nell 1989). The mineral content of
termitaria and the adjacent soils has also been
carefully measured by several researchers (Watson
1977, Steinke and Nell 1989, Holt and Lepage 2000,
Cadet et al. 2004, Masanori and Tooru 2004,
Brossard et al. 2007, Chikuvire et al. 2007). Most
research shows that termitaria contain significantly
higher concentrations of total nitrogen (N) and
exchangeable cations than the surrounding area
(Watson 1977, Steinke and Nell 1989, Jones 1990,
Holt and Lepage 2000, Chikuvire et al. 2007). In
tropical wet–dry climates, downslope erosion could
enhance the soil fertility around termitaria
compared with leached soils away from it (Malaisse
1978). In addition, soil from termitaria has other
positive effects on crops such as suppression of
weeds. For example, Cubitermes soil was found to
suppress Striga infestation on sorghum effectively
in Burkina Faso (Andrianjaka et al. 2007).
Although scientists believe that soil from termitaria
may provide an alternative to chemical fertilizers,
some farmers have resisted leveling termitaria
(Logan et al. 1990). Scientists have expressed
concern about why farmers do not level termitaria
in order to make full use of the land and to facilitate
mechanized tillage operations (Nyamapfene 1986).
Such concerns ignore the spiritual (Geissler 2000,
Copeland 2007) and economic values (Nkunika
1998) that farmers attach to termitaria. The
recommendation of scientists also ignores the fact
leveling termitaria is not sustainable in the long
term. According to Brossard et al. (2007), excessive
use of termitaria soil can affect termite populations
besides mining nutrients.
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Fig. 2. Farmers’ practice of planting crops around termitaria. This farmer in Zomba district of southern
Malawi has planted banana and pigeon pea on an Odentotermes mound.
Farmers’ Termite Management Practices
Although extension programs encourage the use of
pesticides, most farmers in the study sites relied on
indigenous control practices (Table 4). Although
some of these are parts of good agricultural
practices, the lack of critical scientific evaluation
has limited their wider use. Here, we will briefly
discuss the most common farmers’ practices.
Destruction of termitaria and the colony
Farmers in the study sites mentioned different
methods of destroying the colony (Table 4). These
included digging the nest and removing the queen;
burning wood, grass, or cow dung; pouring hot
water, insecticides, rodenticides, or paraffin; and
flooding the nest with rainwater to kill the colony
(Malaret and Ngoru 1989, Nyeko and Olubayo
2005). Although destruction of the colony has been
advocated by researchers (Logan et al. 1990),
success has been limited because of various
constraints including labor requirements and lack
of knowledge about termite biology. This practice
is directed toward mature colonies of the mound-
building species, and species that do not build
mounds (e.g., many Odontotermes and Microtermes
spp.) are often overlooked. Those species that build
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Table 4. Farmers’ indigenous termite management practices in Ethiopia, Kenya, Uganda, Malawi, and
Zambia
Control practices Districts Country (part of
country)
References
Destruction of mound and colony Balaka Malawi (southern) Sileshi et al. (2008)
Katete, Chipata Zambia (eastern) Sileshi et al. (2008)
Tororo Uganda (eastern) Nyeko and Olubayo (2005)
Machakos Kenya (eastern) Malaret and Ngoru (1989)
S Gonder Ethiopia (northern) Jiru (2006)
Plant Euphorbia tirucali in field Balaka Malawi (southern) Sileshi et al. (2008)
Tsangano Mozambique
(northwest)
Sileshi et al. (2008)
Chipata, Katete Zambia (eastern) Sileshi et al. (2008)
Kalomo, Choma, Monze Zambia (southern) Nkunika (1998)
Apply Bobgunnia madagascarensis Chipata, Katete Zambia (eastern) Sileshi et al. (2008)
Kalomo, Choma, Gwembe,
Monze, Livingstone
Zambia (southern) Nkunika (1998)
Spray Tephrosia extract Chipata Zambia (eastern) Sileshi et al. (2008)
Apply wood ash Chipata, Katete Zambia (eastern) Sileshi et al. (2008)
Kalomo, Gwembe, Monze,
Mazabuka, Livingstone
Zambia (southern) Nkunika (1998)
Machakos Kenya (eastern) Malaret and Ngoru (1989)
Tororo Uganda (eastern) Nyeko and Olubayo (2005)
Avoid earthing up (banking) Chiosha Malawi (central) Sileshi et al. (2008)
Chipata Zambia (eastern) Sileshi et al. (2008)
Minimum weeding Chipata Zambia (eastern) Sileshi et al. (2008)
Clean weeding Chipata Zambia (eastern) Sileshi et al. (2008)
Machakos Kenya (eastern) Malaret and Ngoru (1989)
Apply pork, meat, fat to attract ants Mapanje Mozambique
(northwest)
Sileshi et al. (2008)
Machakos Kenya (eastern) Malaret and Ngoru (1989)
Apply cow dung or urine Monze Zambia (southern) Nkunika (1998)
Machakos Kenya (eastern) Malaret and Ngoru (1989)
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mounds are subterranean for the first few years.
Even if mature colonies are killed, the immature
colonies could spread to take over the area (Logan
et al. 1990). In addition, many farmers resist
destroying the mounds (Nyamapfene 1986,
Nkunika 1998) even when they are live or very easy
to flatten (see Fig. 3). Mound destruction may not
be acceptable, probably because termitaria are
sacred places among many communities in Africa
(Geissler 2000, Mguni 2006, Copeland 2007).
Plant materials
Many plant species have been used by farmers
across Sub-Saharan Africa to control termites
(Logan et al. 1990, Nkunika 1998). Among the plant
species mentioned frequently in the specific study
sites and the literature, Euphorbia tirucalli ranks
first. Farmers in Malawi and Zambia believe
planting E. tirucalli in crop fields or applying its
branches in planting holes deters termites (Orr and
Ritchie 2004, Sileshi et al. 2008b). In Tanzania, the
leaves and roots of E. tirucalli are soaked in water
and the solution is sprayed to protect seedlings from
termites (Logan et al. 1990). In Zambia, farmers
apply crushed pods of Bobgunnia (Swartzia)
madagascareinsis in planting holes (Nkunika 1998,
Sileshi et al. 2008b). Extracts from leaves of
Tephrosia vogelii are also used to protect tree
seedlings in Malawi and Zambia (Nkunika 1998,
Sileshi et al. 2008b). The limitation of plant
materials is that farmers’ recipes vary widely. The
mechanism by which these concoctions reduce
termite damage is as yet unclear. Most plant
materials also break down rapidly in the soil and do
not give prolonged protection from termite attack
(Logan et al. 1990). In addition, the hazard they
present to humans and the environment is often
unknown. Therefore, greater care is required in their
use. Rigorous toxicological, safety, and environmental
evaluation is also needed for their wider application.
Wood ash
Wood ash has been widely mentioned as one of the
control practices in eastern and southern Zambia
(Nkunika 1998, Sileshi et al. 2008b) and Nigeria
(Banjo et al. 2003). Logan et al. (1990) summarize
reports about the use of wood ash for termite control.
However, the mechanism by which ash provides
protection against termites is unclear. Variations
also exist on the effectiveness of ash (Nkunika
1998). This information gap demands better
evaluation of wood ash against the most serious
termites for the particular area.
Protein- or sugar-based products
In Uganda, farmers use dead animals, meat, and
sugarcane husks to “poison” Macrotermes mounds
(Sekamatte et al. 2001). Farmers in Tsangano
district of Mozambique mentioned that they used
leftover pork or beef to control termites (Sileshi et
al. 2008b). Similarly, Nigerian farmers bury dead
animals or fish viscera to reduce termite attack on
crops (Logan et al. 1990). In South Africa, Riekert
and van den Berg (2003) experimentally
demonstrated significant reduction in termite
damage to maize using fish meal. Until recently, the
rationale behind this practice had not been clear
(Logan et al. 1990). Sekamatte et al. (2001)
demonstrated that the reduction in termite damage
in plots that received fish meal is due to increased
activity of ants. The protein-based baits resulted in
greater ant nesting near maize plants and reduction
in termite damage (Logan et al. 1990).
Cow dung and urine
Cow dung and urine have been used for termite
control by farmers in the study areas (Malaret and
Ngoru 1989, Nkunika 1998) as well as elsewhere
(Banjo et al. 2003, Nwilene et al. 2008). In
Machakos district of Kenya, farmers smear cow
dung on posts to protect them from termite attack
(Malaret and Ngoru 1989). In Monze district of
Zambia, farmers used fresh cow dung to reduce
termite damage to maize (Nkunika 1998). Similarly,
farmers in southwestern Nigeria believe that goat
and cow dung reduce termite damage (Banjo et al.
2003). Reduction in termite damage to rangeland
using cow dung has been demonstrated in an
experiment conducted in the “Cattle Corridor” of
Uganda (Tenywa 2008). Further research should
establish the effectiveness and the mechanism by
which cow dung reduces termite damage to crops
or trees.
Intercropping
According to farmers in parts of northern Uganda,
maize intercropped with sorghum usually suffers
less termite damage than pure stands. Farmers in
this region regarded the sorghum plants as
reservoirs of the predatory ants (Sekamatte et al.
2003). Intercropping maize with food legumes has
been demonstrated to increase nesting of predatory
ants and reduce termite attack on maize (Sekamatte
et al. 2003). However, the impact of intercropping
on termite damage depended largely on the legume
species in the intercrop (Sekamatte et al. 2003,
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Fig. 3. Farmers’ usually leave termitaria intact when cultivating crop fields. This figure shows live
Macrotermes mounds left in a field ready for planting maize in Balaka district of southern Malawi.
Sileshi et al. 2005). In Uganda, termite attack was
lower in maize–soybean (Glycine max) than in
maize–groundnut or maize–bean intercrops
(Sekamatte et al. 2003). Similarly, in Zambia, some
legume trees in agroforestry reduced termite
damage to maize more than others (Sileshi et al.
2005). These differences could be attributed to
variations in leaf litter, which serves as an
alternative source of food for termites and the thick
canopy in the intercrops that encouraged predation
(Sekamatte et al. 2003, Sileshi et al. 2005).
Weeding and tillage practices
In Malawi and eastern Zambia, farmers avoided
ridging the soil when weeding or reduced the
weeding to a minimum in order to reduce termite
damage on maize crops (Sileshi et al. 2008b).
Tillage and weeding may have negative effects on
termite activity because of the physical disruption
of their feeding galleries, alteration of soil
environment and food resources, and exposure to
predators (Logan et al. 1990, Black and Okwakol
1997). However, conflicting reports exist on the
effects of weeding on termite populations. This
highlights the need for site-specific studies and
recommendations.
Potential Impacts of Disruptive Termite
Control Practices
Until recently, organochlorines, which are regarded
as persistent organic pollutants (POPs) have been
widely used for termite control (Logan et al. 1990,
Langewald et al. 2003). With the banning of POPs,
the search for alternative insecticides has increased.
The most recent development in termite control
involves fungicides targeted at the Termitomyces
spp. (Rouland-Lefevre and Mora 2002). Such
control practices are usually initiated on anecdotal
information rather than on sound scientific inquiry
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into the biology of the local termite species or their
true impact on crops or trees in Africa. None of the
entomological literature in Africa provides any
empirical data on the social, ecological, or economic
risks and trade-offs of control practices. The impacts
of chemical control on human welfare and the
environment have also been largely ignored or
perceived as localized problems. Termitaria are
often the major source of apprehension and targets
for application of pesticides and other control
measures. Termite control using pesticides is likely
to have negative impacts on human welfare and the
environment in at least three ways. First, direct
exposure of farm families to pesticides could occur
because people who apply pesticides usually do not
take precautions or wear protective clothing.
Second, people who consume termites and
mushrooms from treated termitaria could be
exposed to pesticide residues. Third, children and
women can be exposed to pesticides through
consumption of soil from treated termitaria. In
addition, termite control practices could pose risks
to non-target organisms that inhabit termitaria or
consume the soil. In the following sections we will
briefly discuss the categories of organisms briefly.
Non-target termites: Many termite species that are
not mound builders themselves make their nest
within the fabric of existing termitaria (Coaton
1962, Glover 1967, Pomeroy et al. 1991). For
example, in Zambia, Coaton (1962) found more
than 20 different genera inhabiting mounds of other
termites (on average, 3.4 genera per mound). Some
of these species, which are harmless, may be killed
by control practices targeted at the termitaria.
Symbionts: Termite symbioses are characterized by
obligate nutritional mutualisms between the insect
host and various micro-organisms including
bacteria, protozoa, and fungi (Bignell 2000). The
gut of termites is a rich reservoir of novel micro-
organisms (Mackenzie et al. 2007) and biochemical
pathways that may even provide the means to
produce green biofuels from wood (Rubin 2008).
Microbial communities that inhabit termite nests are
also of great genetic and functional diversity that
could be used for phytoremediation (Duponnois et
al. 2006). Among the fungi, members of the genus
Termitomyces live in a mutualistic association with
the Macrotermitinae (Aanen and Eggleton 2005)
and produce edible mushrooms. Termite control
using fungicides targeted at the Termitomyces spp.
(Rouland-Lefevre and Mora 2002) may have
negative impacts on the ecosystem services of
termites, including production of edible mushrooms.
Termitophilous and termitolestic species: Termitaria
also provide habitat for a variety of termitophilous
animals, which live in termite nests and steal or
scavenge food from termites. These include
obligatory inquilines of Insecta, Isopoda,
Collembola, Thysanura, and Psocoptera (Glover
1967, Kistner 1990, Pomeroy et al. 1991).
Termitolestic species live in or near the nest and
regularly prey on the termites (de Visser et al. 2008).
This includes ants, spiders, centipedes, and assassin
bugs (Pomeroy et al. 1991). Many of these
invertebrates are poorly studied and their
community structure and function not known.
Vertebrates that feed and nest in termitaria:
Termites form the major dietary component of many
animals, including invertebrates and vertebrates.
For some animals, such as the aardvark, termites are
the main food (Pomeroy et al. 1991, Peveling et al.
2003). Termites are also the most nutritionally
important insects in the diet of chimpanzees and
gorillas (Tutin and Fernandez 1983, Cipolleta et al.
2007, Deblauwe and Jenssens 2007). During
swarming, termites form a major source of food for
many amphibians, reptiles, birds, and mammals
(Dial and Vaughan 1987). In addition, many
animals including birds, reptiles (e.g., geckoes,
lizards, and snakes), and mammals nest in termitaria
(Glover 1967, Pomeroy et al. 1991). For example,
barbets (Trachyphonus spp.) preferentially nest in
burrows in the wall of termitaria (Pomeroy et al.
1991). The Nile monitor (Varanus niloticus) buries
its eggs in termitaria for incubation (Bennett 1995).
Mongooses are largely dependent on termitaria for
safe den sites (Rasa 1985). Among the modern
insecticides used for termite control, carbamates are
regarded as not toxic to mammals. However, they
are extremely toxic to birds. Therefore, use of
carbamates can kill birds that have eaten poisoned
termites or nest in termitaria. Poisoning the
termitaria is also likely to have negative impacts on
the other organisms.
Animals that ingest soil: Termitaria also serve as
salt licks for both domestic and wild animals. For
example, elephants (Loxodonta africana) are
known to excavate termitaria and eat the soil to
satisfy their need for micronutrients such as iodine,
cobalt, and selenium (Milewski and Diamond
2000). Consumption of soil from termitaria is also
very common among primates (Klein et al. 2008).
Herbivorous animals: Termitaria usually act as
grazing and browsing hotspots for herbivores
(Holdo and McDowell 2004, Loveridge and Moe
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2004, Mobæk et al. 2005). In Uganda, ungulates
grazed preferentially on termitaria compared with
the adjacent savannah (Mobæk et al. 2005). In
Zimbabwe, trees growing on termitaria were
subjected to more intense feeding by elephants and
rhinoceros (Diceros bicornis, Ceratotherium
simum) than trees in the surrounding vegetation
matrix (Holdo and McDowell 2004, Loveridge and
Moe 2004). Therefore, indiscriminate use of
pesticides on termitaria may put the health of
animals that either graze or browse the vegetation
or consume the soil.
CONCLUSION
The review of the general literature and specific
studies leads to the conclusion that local
communities have comprehensive indigenous
knowledge of termite ecology and taxonomy and
apply various indigenous control practices. Farmer
innovation was evident in the diversity of
indigenous termite control practices. Farmers have
a more tolerant attitude toward termites and termite
mounds than do researchers. It is not surprising that
traditional agriculture has coexisted with termites
in Africa.
The review above indicates that termite control
practices can have negative impacts on human
welfare and the environment in a number of ways.
Therefore, more balanced termite management
practices are needed to ensure human welfare and
maintenance of the ecosystem services provided by
termites. Farmers’ indigenous knowledge may help
in improving the existing practices or searching for
environmentally friendly and socially acceptable
management approaches. Farmers’ indigenous
practices depend on intimate knowledge of the local
situation. Unfortunately, indigenous practices have
been dismissed by some researchers as
unsatisfactory without adequate research. This is
because the performance of indigenous practices is
often judged against insecticidal control, which
often gives an immediate result. Instead of
dismissing such practices as ineffective, their
rationale and shortcomings need to be identified to
generate contextual and site-specific knowledge. In
this way, limitations of indigenous practices can be
removed and solutions with local relevance may be
found. This emphasizes the point that ethno-
ecological knowledge is best employed as a
complement to, rather than a substitute for,
scientific knowledge. There are many ways in which
indigenous practices may be fine tuned if farmers
and scientists work together and learn from each
other. Toward that end, we recommend a farmer-
targeted participatory research approach to build
coherent principles for termite management in
Africa.
Responses to this article can be read online at:
http://www.ecologyandsociety.org/vol14/iss1/art48/
responses/
Acknowledgments:
The field work by the first three authors was
supported by the Sustainable Agriculture Initiative
of the Council for the Development of Social Science
Research in Africa (CODESRIA) and International
Foundation for Science (IFS). Financial support for
part of the field work and manuscript preparation
also came from the United States Agency for
International Development (USAID), Canadian
International Development Agency (CIDA),
Swedish International Development Agency (Sida),
and Irish Aid through the World Agroforestry Centre
(ICRAF).
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