Termite Management in Tropical Agroforestry Termite Management in Tropical Agroforestry

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Termite Management in Tropical
Agroforestry

Termite Management in Tropical
Agroforestry
P.O.Y. Nkunika, Entomologist, University of Zambia
W.G. Sileshi, Agroecologist, World Agroforestry Centre
P. Nyeko, Entomologist, Makerere University
B.M. Ahmed, Entomologist, University of Melbourne

Copyright © Authors 2013
All rights reserved. No part of this publication may be reproduced,
stored in a retrieval system or transmitted in any form or by any means
without the prior permission of the copyright holder(s)
ISBN: 978-9982-03-067-0
Published by the University of Zambia Press
on behalf of the Authors
P.O. Box 32379
Lusaka 10101
Zambia

CONTENTS
Page
FOREWORD............................................................................................. vi
ACKNOWLEDGEMENTS....................................................................... vii
PURPOSE OF THIS BOOK...................................................................... viii
CHAPTER ONE........................................................................................ 1
INTRODUCTION..................................................................................... 1
1.1 Agroforestry Practices.................................................................. 1
1.1.1 Multi-strata Agroforestry..................................................... 2
1.1.2 Agroforestry Parklands........................................................ 3
1.1.3 Silvopastoral Systems.......................................................... 3
1.1.4 Alley Cropping..................................................................... 3
1.1.5 Improved fallows and Intercropping................................... 4
1.1.6 Woodlots and Rotational Woodlots..................................... 4
1.1.7 Relay Cropping.................................................................... 5
1.1.8 Conservation Agriculture with trees.................................... 5
1.2 Advantages of Agroforestry.......................................................... 5
CHAPTER TWO....................................................................................... 9
TERMITE BIOLOGY AND BEHAVIOUR.............................................. 9
2.1 The Structure and Social Organisation of Termites...................... 9
2.1.1 Reproductives (Alates)........................................................ 10
2.1.2 Workers................................................................................ 11
2.1.3 Soldiers................................................................................ 12
2.2 Life Cycle of Termites.................................................................. 13
2.3 Termite Nests................................................................................ 14
2.4 TermiteClassicationandIdentication...................................... 16
2.5 Termite Food................................................................................ 19
CHAPTER THREE................................................................................... 21
BENEFICIAL EFFECTS OF TERMITES................................................ 21
3.1 LivelihoodBenets....................................................................... 21
3.1.1 Termites as Human Food..................................................... 21
3.1.2 Termite Mounds as Sources of Mushrooms........................ 22
3.1.3 Termites Improve Agricultural Land................................... 23
3.2 Provision of Ecosystem Services.................................................. 23
3.2.1 Termites as Food for Animals.............................................. 23
3.2.2 Termite Mounds as Sources of Minerals............................. 24
3.2.3 Termite Mounds are Hotspots of Livestock Feeding........... 24

3.2.4 Termites help Regenerate Degraded Land and Tree
Establishment....................................................................... 24
3.2.5 Termite Mounds are Hotspots of Biodiversity..................... 25
CHAPTER FOUR..................................................................................... 27
TERMITES AS PESTS............................................................................. 27
CHAPTER FIVE....................................................................................... 31
TERMITE MANAGEMENT..................................................................... 31
5.1 General Principles for Reducing Termite Damage....................... 32
 5.2 SpecicControlMethods............................................................. 36
5.2.1 Chemical Control................................................................. 36
5.2.1.1 Alternative Control Options..................................... 36
5.2.1.2 Microbial Preparations............................................. 38
5.2.1.3 Wood Ash................................................................. 38
5.2.1.4 Cow Dung and Urine............................................... 38
5.2.1.4 Destruction of Mounds and Colonies...................... 39
CHAPTER SIX.......................................................................................... 41
CONCLUSION.......................................................................................... 41
REFERENCES.......................................................................................... 42
APPENDICES........................................................................................... 45
Appendix 1.................................................................................... 45
Appendix 2.................................................................................... 49
Appendix 3.................................................................................... 52

vii
FOREWORD
This book is a synopsis of termite biology, ecology and its management
in agroforestry, which is an integration of crop production, livestock and
forestry practices, on the same piece of land. Indeed, trees are one of the
most promising means known to better adapt farming systems to climate
change, and to act as sinks for absorbing carbon dioxide. This book shall
assistresearchers,students,farmers, extension ofcers, policymakersand
Non-Government Organisations (NGOs) to understand the signicance of
termites in agroforestry.
Although termites are usually considered pests, some are not pests at
all. While the pestiferous species have negative aspects, they can provide
manybenets.Forexample,termitesloosen,aerateandenrichsoilthrough
tunnelling, litter decomposition and recycling of soil organic matter in
conservation farming. Some edible mushrooms such as Termitomyces grow
mainly around termite mounds. Some termite species are also an important
source of proteins to human beings, animals and poultry.
The University of Zambia is appreciative of the initiative by the authors
to undertake this project, which will contribute to the body of knowledge of
thisnewandexcitingeldoftermitemanagementinagroforestry.Sincethe
initiation of the project in 2004 by the Department of Biological Sciences,
of the School of Natural Sciences, its impact and signicance have been
acknowledged by researchers not only in Zambia, Malawi and Uganda but
asfaraeldasAustralia.
It is my hope that this publication will inspire researchers and other scholars
to do further research in this much neglected, but very important area.
Professor S.F. Banda
Dean, School of Natural Sciences

viii
ACKNOWLEDGEMENTS
The authors acknowledge Mr Christopher Phiri of Crop Science Department,
School of Agricultural Sciences, University of Zambia for reviewing the initial
manuscript and providing useful suggestions. We appreciate the contribution of
the Dean of the School of Natural Sciences, University of Zambia, Prof. Samuel
F. Banda for his logistical support and encouragement, which kept our spirits
high.
WeappreciatethenancialsupportreceivedfromtheSustainableAgriculture
Initiative of CODESRIA-IFS in 2003-2005. The second author also gratefully
acknowledgesnancialsupportfrom the Canadian InternationalDevelopment
Agency (CIDA), Swedish International Development Agency (SIDA) and Irish
Aid.
The authors thank Mr Gerald Mwale of the Department of Mass
Communication, Mrs Miriam J.K. Nkunika and Mr Billy Nkunika for their
contributions during the drafting of the initial manuscript. We also thank
Professor Joseph Moughalu of the University of Ife who reviewed the entire
manuscript and gave constructive comments. We are also grateful to several
anonymousreviewersforhelpingustoimprovethenaldraft.

ix
PURPOSE OF THIS BOOK
This book is intended to be used as a guide to management of termites in
agroforestry and farmsteads where termites pose serious problems. Emphasis
isplaced ontropicalagroforestryandmorespecicallyinAfrica. However,it
is also hoped that the book will be of use in other continents where termites
pose problems. It has been written in response to the growing need to improve
understanding of termite problems and taking appropriate action based on such
knowledge. In the past, blanket recommendations for termite control have
resultedininefcientandunsustainableuseofresourcesandhavehadnegative
impacts on the environment. Control practices have often been initiated on
anecdotalinformationrather thanonsoundscientic inquiryintothebiology
and ecology of the local termite species and their true impact on crops or trees.
It is important to note that there is no single method that can be universally
recommended for termite control because several species may be found in an
areaandeachspecieshasauniquebiologyandecology.Therefore,thelevelof
control depends on the knowledge of the termite species, the tree or crop species,
climatic conditions and other environmental factors. Our aim is to promote a more
sustainable management of termites, i.e., an approach 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) utilisation of termites and associated resources without exhausting them.
Thiscanbeachievedthrough adequate understanding of termite biologyand
application of control measures based on ecological principles.
The intended users of this manual are farmers (subsistence as well as
commercial),extensionofcers,researchers,pestcontroloperatorsandstudents.
The aim is to provide basic information on termites and help people to take
more informed actions to manage termites in agroforestry.
This book is presented in six chapters. Chapter one introduces the concept of
agroforestry, agroforestry practices, different types of agroforestry, advantages
of agroforestry and introduces the subject of termite management in agroforestry.
Chapter two provides a description of the biology of termites including their social
organisation,lifecycle,nestingbehaviour,classicationandfood.Chapterthree
providesasummaryofthebenecialaspectsoftermites.Chapterfourprovides
a brief discussion on common pestiferous termites and the damage they cause
tocropsandtrees.Chapter vefocuseson principlesthatshould befollowed
for sustainable management of pestiferous termites in agroforestry with less
emphasis on control and Chapter six ends with concluding remarks and prods
scholars to do further research in this much neglected, but very important area.

1
Chapter One
INTRODUCTION
There are many traditional land use practices that involve production of trees,
agricultural crops and livestock on the same piece of land. These land use systems
are collectively called agroforestry, which is a new name for old practices. In
most cases, the major objective of these practices is not tree production but
production of food and other goods and services. Today, woody plants continue
to supply farmers with a number of valuable products such as fruits, fuel wood,
timber,fodder,breandmedicines.Trees,inparticularnitrogen-xingspecies,
also add fertility to soil, thereby increasing yields of agricultural crops. The trees
may also be deliberately retained for cultural or religious reasons.
Agroforestry is a concept of integrated land use that combines elements of
agriculture and forestry in a sustainable production system. Agroforestry is
broadlydenedasthesetoflandusepracticesinvolvingadeliberatecombination
of trees (including shrubs, palms and bamboos) and agricultural crops and/or
animals on the same land management unit in some form of spatial arrangement
or temporal sequence such that there are signicant ecological and economic
interactions between tree and agricultural components (Sinclair, 1999). In this
denition, agroforests, which are complex agroforestry systems looking like
and functioning as natural forest ecosystems, are integrated into agricultural
managementsystems.Theabovedenitionispreferredtotraditionaldenitions
because farmers and forest dwellers, where agroforestry has developed as a
signicantlandusepractice,havetendedtopracticeagroforestryusingvarious
methods. For instance, either by integrating many tree species in various
productive niches on their farms or by managing biodiverse forest resources.
Therefore, agroforestry is an approach to land use involving a deliberate and
purposeful combination of trees with crops and/or animals.
1.1 Agroforestry Practices
An agroforestry practice refers to the distinctive arrangement of woody species
(i.e. trees and shrubs), crops and livestock in space and time. In terms of function,
the woody components of agroforestry may be categorised as follows:
1. Fertiliser trees: This refers to all nitrogen-xing leguminous or non-
leguminous woody perennials deliberately planted for soil fertility
improvement in arable land, pastures or other land use systems. The
species may be trees or shrubs that x nitrogen through associations
with symbiotic bacteria such as Rhizobia and Frankia. The contribution

2
of fertiliser trees to soil fertility mainly comes from nitrogen inputs via
deepcaptureandbiologicalnitrogenxation.Inaddition,fertilisertrees
canprovideadditional products(e.g.,rewood,poles, seeds,etc.)and
services (e.g., soil and water conservation).
2. Fruit trees: perennials that produce edible fruits may be planted in blocks,
scattered on farmland, alongeldboundaryoraroundthehomestead.
3. Fodder trees: perennial leguminous trees and shrubs that provide
protein sources for livestock can supplement pasture and also provide
an effective insurance against seasonal feed shortages or during time
of drought. Trees may be planted in blocks, along soil conservation and
contour bands, eldboundaryoraroundthehomestead.
4. Timber trees:perennialsspecicallyplantedforproductionoftimberin
a variety of spatial arrangements. However, timber may be a secondary
product, harvested only after the tree has served its primary production
or service role (e.g., fertilising the soil, controlling erosion and producing
fruits, vegetables, fodder, medicines, resins, shade, etc.)
In terms of spatial arrangement, agroforestry practices may take the form of
dense mixed stands (as in multi-strata home gardens), sparsely mixed stands
(as in many parklands and silvopastorial systems) or strips of varying width
(as in alley cropping, boundary planting, contour planting, and intercropping).
Temporal arrangements of plants in agroforestry may take the form of shifting
cultivation, improved fallow or some silvopastorial systems that involve rotation
of grass leys with woody species. The common agroforestrypracticesarebriey
described below.
1.1.1 Multi-strata Agroforestry
The term multi-strata agroforestry comprises all tree crop-based land use
systems that have two or more vegetation layers ranging from plantations of
coffee (Coffea spp.), cacao (Theobroma cacao), or tea (Camellia sinensis) mixed
withamonospecicoverstoreyofshadetreestohighlydiversiedsystemssuch
as home gardens and ‘agroforests’ (Nair et al., 2008). In the humid tropics, trees
in the genera Albizia, Acacia, Entada, Erythrina, Gliricidia, Inga, Leucaena
and Millettia are among the commonly used trees in cacao and coffee growing
countries. Cacao has been grown under gliricidia (Gliricidia sepium) shade, and
hence the Spanish name for gliricidia is ‘madre de cacao’, literally meaning
‘mother of cacao’. Cardamom (Amomum subulatum), the most important
perennial cash crop in the eastern Himalayas is cultivated predominantly under
Alnus nepalensis. In Africa, multi-strata agroforestry is represented by the tree-
crop farming system and highland perennials farming system where cacao,

3
coffee, palm oil, rubber, yams, cassava, banana, plantain, enset, coffee, sweet
potato, beans, maize and livestock are an integral part. Depending on the species,
the shade tree can be regularly pruned for soil improvement or left to grow fully
toproducerewoodandtimber.
1.1.2 Agroforestry Parklands
Parklandsconsistofscatteredtrees occurring on cultivated land, fallowelds
or pasture, and are widespread in the semi-arid tropics of Africa and Asia. In
the semi-arid areas of the West African Sahel and parts of Eastern and Southern
Africa, the best known parklands are those involving faidherbia (Faidherbia
albida), Acacia spp., Parkia biglobosa and Vitellaria paradoxa. The trees are
derived from natural regeneration, but protected and managed by farmers. In
some countries such as Mali, parklands occupy 90 per cent of the agricultural land
and support livelihoods of over 2.5 million people (Boffa, 1999). Crops such as
sorghum,millet,groundnuts,sesameandlivestockformasignicantcomponent
of these farming systems and hence they represent an agrosilvopastoral system.
1.1.3 Silvopastoral Systems
Silvopastoralsystemsmaybedenedasanagroforestrypracticethatintegrates
trees with animal production. Silvopastoral systems may be divided into two
broad categories: tree fodder (hereinafter called protein banks) and grazing
systems. In the protein bank approach, the animals are stall-fed with fodder from
trees or shrubs grown in blocks on farms (Nair et al., 2008). The usual practice
is cut-and-carry management, which involves removing tree prunings and cut
grassfrom theeldandfeedingstabled animals.Ingrazingsystems,livestock
are allowed to graze on pasture under widely spaced or scattered trees. In the
moreextensivegrazingareas,N-xing trees are increasingly being plantedin
association with improved grasses to increase carrying capacity and productivity
of grazing cattle. In the more intensively managed areas, trees such as Leucaena
and Gliricidia are planted in pasture or protein banks.
1.1.4 Alley Cropping
Alleycropping isdenedasanagroforestrypracticewhere treesorshrubsare
grown in wide rows and arable crops (cereals, legumes or horticultural) or pasture
grasses are cultivated in the alleys between the tree rows. This allows integration
of trees into conventional agricultural sites for simultaneous production of
crops and woody biomass which enables farmers to diversify the production of
marketable goods. Therefore, it is one of the emerging agroforestry practices
globally. In the temperate areas, alley cropping of high value (e.g., timber, fruit

4
or nut-bearing) trees with crop or pasture is one of the more common agroforestry
practices. In the tropics, alley cropping was developed as an alternative to slash-
and-burn agriculture or for erosion control on sloping land, where it is sometimes
called hedgerow intercropping (Kang et al., 1990). In the humid and sub-humid
tropics of Africa, alley cropping is practised in the cereal-root crop mixed
and maize-mixed farming systems where it involves growing maize, beans or
cassava between rows of perennial woody legumes in the genera Acacia, Albizia,
Calliandra, Flemingia, Grilicidia, Inga, and Leucaena and Sesbania. The woody
species may be coppiced and the trees are periodically pruned and their biomass
is applied either as mulch or incorporated into the soil.
1.1.5 Improved Fallows and Intercropping
Improved fallows involve the rotation of crops with woody species for the rapid
replenishment of soil fertility on crop land (Sanchez, 1999). This is an alternative
to natural vegetation fallows, which take longer to restore soil fertility. A piece
oflandisplantedwithfast-growingN-xingtreesorshrubsfor2-3years.Crops
are planted after cutting-back the woody species. Tree species used in improved
fallows can be either non-coppicing or coppicing. Non-coppicing species used
in improved fallows include Sesbania spp., Tephrosia spp. and pigeon pea, and
these do not regrow when cut at the end of the fallow period, typically after 2-3
years of growth. Therefore, after 2-3 years of cropping these species have to
be replanted. This constitutes rotational fallowing. Coppicing species include
members of the genera Gliricidia, Acacia, Leucaena, Calliandra and Flemingia,
which do not need replanting. Crops are planted between the rows of stumps,
and this can be managed by periodic pruning in an intercropping arrangement. A
typical example is gliricidia-maize intercropping in Malawi and Zambia, which
has been demonstrated to achieve stable increases in crop yields over several
years without the need to invest in inorganic fertiliser (Sileshi et al., 2012).
1.1.6 Woodlots and Rotational Woodlots
Woodlots are usually planted in areas not suitable for crop production. Trees
can be managed for supply of timber, fuelwood, construction materials and
production of non-wood products (e.g., caterpillars, natural dyes, gum, wax and
honey). There are a variety of options for planting woodlots. Examples include
block planting, boundary planting, contour planting, homestead planting and
scattered trees on farm land. The rotational woodlot is a variant of improved
fallows,wherefoodcropsareintercroppedwithleguminoustreesduringtherst
2-3years,andthenthetreesarelefttogrow,harvestedinaboutthefthyear,and
food crops are replanted.

5
1.1.7 Relay Cropping
Relay cropping is the planting of herbaceous green manure legumes or fast-
growing shrubs and trees between rows of an already established crop.
The legumes continue to grow after the crop has been harvested. As farmers
prepare land for the next season, they clear-cut the legume and incorporate the
biomass into the soil. Trees in the genus Sesbania, Tephrosia and pigeon pea
are recommended for relay cropping. This works well on small farms, and the
benetcanbeseenimmediatelyafteroneseasonoftreegrowth.
1.1.8 Conservation Agriculture with Trees
The integration of conservation agriculture practices with agroforestry practices is
called conservation agriculture with trees. This practice isbasedonveimportant
principles; (1) minimising soil disturbance, (2) maintaining crop residues for
soil cover, (3) crop rotation, (4) good agronomic management practices, and (5)
incorporatingnitrogen xingtrees andshrubstoensurea more permanent soil
cover and increase soil organic matter.
1.2 Advantages of Agroforestry
Agroforestry ensures a multifunctional agriculture that provides food (fruits, crop
yield), saleable products (e.g., fruits, timber, fodder, medicinal and pesticidal)
and services such as shade, soil improvement and watershed management. The
advantagesincludelaboursaving,livelihoodstrengtheninganddiversication.
Labour Saving
Many agroforestrypracticesimprove the efciencyoflabourthrough positive
interaction between trees and farm enterprises (FAO, 2007). For example,
woodlotssavetimeandlabourforfetchingrewoodbyincreasingsuppliesand
bymovingthesourceofrewoodclosertohome.
Livelihood Strengthening
The presence of trees often increases security of land tenure. Planting trees
reduces environmental degradation while producing by-products like food, fruit,
rewoodandfodderthatstrengthenlivelihoods.
Livelihood Diversication
Oncedemandforwoodinthecommunityhasbeensatised,polesandrewood
may be developed as a source of income. Some farmers may also raise tree
seedlings or produce seeds for sale.

6
Agroforestry also offers opportunities for the development of climate-smart
agriculture; agriculture that sustainably increases productivity (food security),
resilience (climate change adaptation) and reduces greenhouse gas emissions
(mitigation) (FAO, 2012).
Despite the enormous social and ecological benets, agroforestry practices
had largely been under-utilised until the formation of the International Centre
for Research in Agroforestry (ICRAF) now branded as the World Agroforestry
Centre. ICRAF continues to be the global leader in agroforestry research. Recently,
agroforestry practices have also gained currency in research, academic and policy
circles with the emergency of the concept of sustainability. Several global and
regional networks and associations of professionals now spearhead agroforestry
education, research and development. The African Network for Agriculture,
Agroforestry and Natural Resources Education (ANAFE) was launched in
1993 by seventeen universities and twelve technical colleges in sub-Saharan
Africa offering education programmes in agroforestry, agriculture, forestry and
natural resource management. In the USA, communication and/or technology
transfer at the federal level has been enhanced through the establishment of the
joint inter-agency USDA FS/NRCS National Agroforestry Centre in Lincoln,
Nebraska. Scientists, practitioners and landowners formed the Association for
Temperate Agroforestry (AFTA) in 1991. AFTA, based at the University of
Missouri Centre for Agroforestry is now a very large network with membership
in the US, Canada and overseas. The European Federation of Agroforestry
(EURAF) has membership from seventeen European countries. Several major
global agroforestry congresses have also been held, where researchers shared
ndings.TheAFTAhassofarheldtwelvebiennialconferencesonagroforestry.
Atthegloballevel,twocongressesweresuccessfullyconducted.TherstWorld
Congress of Agroforestry, held on 27 June 2004 in Florida was attended by close
to 500 participants from over eighty countries and 750 presentations were made.
The second World Congress of Agroforestry held from 23-28 August 2009 in
Nairobi,Kenyaattracted1200participantsfrom ninety-sixcountries.Therst
Europeanscientic conferenceonagroforestryorganisedbyEURAFwasheld
in Brussels from 9-10 October 2012, attracting participants from seventeen
(17) European countries and more than fty papers were presented. During
the EURAF conference, an event titled ‘Agroforestry: Trees for a Sustainable
European Agriculture’ was also organised at the European Parliament.
Policy support for agroforestry is also increasing. For example, in the
European Union (EU), the Rural Development Regulation 1698/2005 (2007-
2013) supports agroforestry. In the USA, over twenty states have legislation that
pertains either directly or indirectly to agroforestry (Cutter et al., 1999). Direct
legislation supporting agroforestry is available in Hawaii, Iowa, Maryland,

7
Missouri, Minnesota, Nebraska, South Dakota and Virginia. The states of
Delaware, Illinois, Michigan, Mississippi, New Jersey, New Mexico, New York,
North Dakota, Utah, Washington and Wisconsin have indirect effect legislation.
Under direct legislation, most states provide some type of cost-sharing for
approved practices while tax reduction and cost-sharing were favoured in states
with indirect legislation (Cutter et al., 1999).
As a result of all these developments, agroforestry is receiving long-overdue
attention as a resource efcient, eco-agricultural approach to farming. Many
government institutions, NGOs, grassroot and community based organisations
are widely promoting agroforestry for a variety of reasons. Widespread adoption
of agroforestry means intensied cultivation of trees, in time and space. This
may bring about a change in the landscape, pest and disease composition. One
of the major pest problems in agroforestry in the tropics is pestiferous termites,
which may cause complete destruction of tree seedlings and saplings as well
as associated crops. This makes them more important in agroforestry than
many other pests. Increased termite problems with tree planting, are reported
throughout sub-humid and semi-arid regions. Fruit tree planting in drier areas
might be more sustainable than annual crops because the latter are generally
more susceptible to drought than the former. The aim of this book is to bring to
the fore, basic information on the biology, ecology and practical management
options for termites in agroforestry.

9
Chapter twO
TERMITE BIOLOGY AND BEHAVIOUR
‘.... When we come to consider the order in these insects, and their subterranean
cities, they will appear foremost on the list of the wonders of creation.’
H. Smeathman, 1781
2.1. The Structure and Social Organisation of Termites
Termites are wrongly called ‘white ants’, yet they are totally unrelated to ants.
Termites are classied in the order Isoptera, while ants belong to the order
Hymenoptera. Termites are structurally most closely related to cockroaches
(Blattodea) in that one Australian species of termites, Mastotermes darwiniensis
Froggatt, which has an anal lobe in the hind wing and an egg mass like the ootheca
of cockroaches, is looked upon as a sort of connecting link between the termites
and the cockroaches. Other differences between termites and ants are that, ants
have a constricted (narrow) waist while termites have an unconstricted abdomen
broadly joined to the thorax. Ants have elbowed antennae while termites have
moniliformorliformantennae.
Termites are social insects that live in colonies with a caste system involving
sterile individuals (workers and soldiers) and reproductive individuals (queen
and king). One of the major differences between termites and other social
insects, such as ants, bees and wasps, is that the other insects have larval and
pupal stages that are not active within the colony, while termites do not have
larval and pupal stages. Instead, eggs of termites hatch into workers that are
active within the colony. Another difference between termites and other social
insects is that the male termite called king remains with the female throughout
her lifetime and does not die after mating. Termites are eusocial (‘true social’)
insects.Eusocialityisdenedbythreetraitswhichaptlydeneatermitecolony,
namely:
1. Division of labour, with a caste system.
2. Co-operation among colony members in tending the young.
3. Overlap of generations capableofcontributingtocolonytness.

10
An excellent introduction to the sociality of termites and the relationship
between different castes is given in E.O. Wilson’s, The Insect Societies (1971).
Inthefollowingdiscussion,wewillbrieydescribethecastesysteminatermite
colony, which consists of reproductives, workers and soldiers.
Larvae on fungus comb (Photo: G.W. Sileshi)
2.1.1 Reproductives (Alates)
The primary reproductives (winged adults or alates) are able to y, and are
commonlycalledyingtermites.Thesecomprisemalesandfemales,whosesole
purposeistostartnewcoloniesandbecomethefuturekingandqueenoftheir
new colony. In the genus Macrotermesarefoundediblealates.Thequeenlays
eggs which hatch into larvae that then develop into workers, soldiers or nymph/
developing alates.
Primary reproductives before shedding wings
(Photo: G.W. Sileshi)
Future queen (in foreground) and king (in
background) after shedding their wings
(Photo: G.W. Sileshi)

11
Primary reproductives disperse, mate and found a new colony. Primary
reproductives have rounded heads with large compound eyes and antennae
which have 10-30 segments. In the period between leaving a colony and the
establishment of a new one, these individuals bear two pairs of similar shaped
wings. These have simple venation without cross veins and are shed along a basal
(humeral)vein,leavingbehindfourtriangularstumpsorscales.Thequeencan
sometimes grow up to six centimetres long while the other castes are generally
lessthanonecentimetre.Theroleofthekingistofertilisethequeen’seggs.The
kingcloselyresemblesthequeeninappearancewiththedifferencebetweenthem
beingthe largesize ofthe queen’sabdomenonceitisswollenwitheggs.This
condition is referred to as physogastry. A Coptotermesqueen canproduceone
hundredeggsperdaybutmoreadvancedtermitequeens,suchasMacrotermes,
can produce 30 000-40 000 eggs per day and up to 10 million a year. Fertilisation
of eggs may, in some species, be monthly (Pearce, 1997).
2.1.2 Workers
Workers are soft-bodied with rounded heads, pale white in colour and are blind,
wingless, sterile and about 3-4 mm in length. They comprise more than 80 per
cent of the colony population. They build and repair the colony nest, shelter tubes
andgalleries.Theytendthequeen,theeggsandyounginthenursery,forageand
gather food and feed all the other castes. Workers immediately move away from
the light into their colony system when nests or shelter tubes or galleries are
broken open. With the exception of the diurnal harvester termites (Hodotermes
spp.), these insects are generally unpigmented and lightly sclerotised. Worker
termites have different roles and are often of two or more sizes. Therefore,
Queen of Macrotermes falciger (Photo: P.O.Y.
Nkunika)
Gravidtermitequeen(centre)surroundedby
workers (Photo: G.W. Sileshi)

12
polymorphism, based on age and sex, is common in the worker caste. In the
families, Kalotermitidae, Termopsidae and some Rhinotermitidae, there is no
true worker caste.
Worker tasks are performed by late instar
nymphs or pseudergates. A pseudergate is
a non-reproductive, non-soldier individual
that diverges from the imaginal line at a
relatively late stage through a regressive
or stationary moult (Thorne, 1996). Other
worker functions include grooming of
individuals of the dependent castes, digging
tunnels, locating food and water, maintaining
the colony micro-climate conditions, and
building and repairing the nest. Worker
castes are the ones which cause damage to
crops and trees.
2.1.3 Soldiers
Soldiers have larger heads that are longer and wider than those of the workers.
Soldiers of most species of termites include both males and females. Although
the gonads are present in males, they are undeveloped. The heads are greatly
modied, heavily sclerotised, and greatly enlarged, sometimes exceeding the
size of the rest of the body. Eyes are usually vestigial or absent and the antennae
aremoniliformorliform.
Differing sizes of soldiers are found in
some species. Major and minor soldiers
are common in some Termitinae. In the
Rhinotermitinae and Schedorhinotermes,
there are two distinct types of soldiers that
differ in size, head shape and mandibles
(Pearce, 1997).
In some species, the mandibles are reduced
and the head is modied into a long nose
from which a toxic glandular secretion is
redfromthetip.Duringattack,thesurvival
of the species depends on this caste system.
The soldiers’ job is to defend the colony from
any unwanted animals.
Workers of Macrotermes sp.
(Photo: G.W. Sileshi)
Soldier of Macrotermes sp.
(Photo: G.W. Sileshi)

13
Soldiers that have enlarged jaws, or reduced mandibles, cannot feed themselves
and they have to rely on workers for this. This process of feeding of one colony
member by another is known as trophyllaxis. Some termite species belonging to
the sub-family Apicotermitinae have no soldiers (Sands, 1972).
2.2 Life Cycle of Termites
Termites are hemimetabolous insects, which means that the life cycle lacks
distinctlarvalandpupalstagesas is the case in, butteries and moths,which
are holometabolous. Although details differ depending on the species, in general
the life cycle is as follows: a colony is founded following the release of winged
reproductive individuals (alates) from the parent colony. Depending on the
species involved, alates may be released at irregular intervals over a period of
several weeks, or in some species, all colonies in an area will release their alates
atoneparticulartimeoftheyearandoftenataspecictimeoftheday.Alates
areweakiersandafterashortnuptialight,theydroptothegroundandshed
their wings. These wingless individuals are referred to as de-alates. The female
attracts the male by holding her abdomen up, presumably emitting a pheromone.
Paired males and females then form tandems, which set off in search of a suitable
nesting site in the soil or wood, depending on the species.
Whena suitablesitehas beenlocated,thepairbecomes thequeenandking
of the new colony, burrowing into the substrate and sealing themselves in a
chamber,wheremating takesplace.Thersteggsarelaidshortlyaftermating
andtheroyalpairtendstotheyoungwhentheyemerge.Therstbroodproduces
whitish, soft-bodied immature individuals, which undergo a series of moults and
develop into workers. Oviposition is resumed when the initial brood is able to
maintainthemselvesandfeedthequeen.
Soldiers will then be produced.
Egg production steadily
increases until it is a continuous,
uninterrupted process. Mating
occurs periodically throughout
the life of the royal pair. Workers
and soldiers are produced
initially and only when the
colony has reached certain age
and size, will reproductives be
produced.
Queen
Egg
Larva
Nymph/Developing
Alates
Female and Male
Winged Alates
King
Soldier
Worker
Life Cycle of Termites

14
Mature colonies, which take years to develop, may range in size from a few
hundred individuals, as in the Kalotermitidae to a million or more in some
Termitidae. A mature colony usually has one king and queen, which reign
throughouttheirlives.However,shouldoneof themdie,or ifthequeen stops
laying sufcient eggs, the rest of the colony will detect this, most probably
by means of pheromones. The production of secondary reproductives is then
initiated. These reproductives do not have the same egg-laying capacity as the
queen of the original pair and consequently, several secondary reproductives
may be found in one nest (Thorne, 1984).
2.3 Termite Nests
Like other social insects, termites live in colonies which live in underground
nests or in above ground mounds. For instance, Hodotermes, Microtermes and
Ancistrotermesspeciesbuildundergroundnests,whichcanbedifculttolocate.
Other termites build their nests in wood, for example, the Kalotermitidae, live
in dry wood, and the Termopsidae live in damp wood (Lee and Wood, 1971).
Other termites build mounds above-ground called, termitaria, often containing
many thousands of individuals. Some of the most famous examples of termite
mounds can be found in Africa, where they may tower nine metres (30 feet)
high, contributing to the ecology landscape of the area (Sileshi et al., 2009;
2010). Termite mounds are fascinating from an engineering perspective, as they
involve immense cooperation and ingenuity to build (French & Ahmed, 2011).
The diagrams below show examples of common termite mounds in Africa.
Termite mounds come in all shapes and sizes, ranging from small, soft mounds
near the entrance to the underground nest to huge, ornate structures sometimes
referred to as cathedral mounds. For example, Macrotermes species build
large
above-ground mounds from which they forage outwards for distances
of up to
50 m in runways. Odontotermes species build both subterranean and
above-ground nests. Termites construct shallow subterranean foraging galleries
Cubitermes mound
(Photo: G.W. Sileshi)
Odontotermes mound
(Photo: G.W. Sileshi)

15
radiating from the nest for distances of up to 50m. The main galleries give rise to
a network of smaller galleries from which foraging parties can exploit potential
food sources over extensive areas of land.
Termites build their mounds using soil and saliva and a complex system of
galleriesorcoveredrunwayswhichtheyuseinsearchingforrequisitessuchas
food, moisture and soil particles. Vents within the mound facilitate the regulation
of mound temperature and humidity (Lee and Wood, 1971). A termite nest
typically stretches underground beneath the mound. One of the key features of
a termite mound is temperature regulation, with termites opening and closing
vents to achieve a stable temperature (French & Ahmed, 2010). Mounds are
also used to control humidity, with conditions which can be so stable that the
humidity rarely varies by more than one per cent.
Termites in the sub-family Macrotermitinae cultivate fungus gardens inside
the mound. These termites live in a symbiotic relationship with fungi of the
genus Termitomyces. Termites cultivate the fungi on a special medium called the
fungus comb, made by the termites. The termites continuously supply the fungus
gardens with plant material including wood, leaves or grass. On these combs,
Macrotermes falciger mound
(Photo: P.O.Y. Nkunika)
Macrotermes michaelensis mound
(Photo: G.W. Sileshi)
Fungus comb inside a Macrotermes mound
(Photo: G.W. Sileshi)
Close up of fungus comb and termite larvae
feeding on fungus (spherical white objects)
(Photo: G.W. Sileshi)

16
the fungi form white blobs called fungal heads containing spores, which are
eaten by termites. However, different termite species use the fungi and the comb
in different ways; some eat the fungus for food (and do not eat anything else),
others eat the fungi for their enzymes and eat plant or comb material as well.
2.4 Termite Classication and Identication
Table1givesasummaryoftheclassicationofthemajorgroupsandapproximate
number of species known to occur in Africa, together with their feeding habits.
Termites are usually divided into lower and higher termites. The lower termites
in Africa belong to the families Kalotermitidae, Termopsidae, Rhinotermitidae
and Hodotermitidae. Members of the family Kalotermitidae feed mainly on
drywood,donotconstructdenite nests, and live in small coloniesinsound
or dead wood. Members of the family, Termopsidae feed on and nest within
damp and decaying wood. The family, Rhinotermitidae consists of mainly
subterranean, wood-eating termites. The family, Hodotermitidae consists of
the harvester termites, which are among the most destructive pests of pasture,
crops and structural timber (Uys, 2002). This family is represented by the genus
Hodotermes in East Africa and Hodotermes and Microhodotermes in Southern
Africa. Both species build subterranean nests and feed on grass.
The higher termites all belong to the family Termitidae. This group shows
considerable variation in feeding and nesting habits and social organisation.
Unlike the lower termites, their feeding is not limited to wood; some feed
exclusively on soil, while others ‘cultivate’ and consume fungi. In Africa, the
Termitidae are represented by over 600 species (>90 per cent of all known
species) in the four sub-families (Apicotermitinae, Termitinae, Macrotermitinae
and Nasutitermitinae). In Africa, the sub-family Apicotermitinae currently
Macrotermes bellicosus mound
(Photo: P. Nyeko)
Macrotermes subhyalinus mound dug to
removethequeen(Photo:P.Nyeko)

17
consists of seventy species, while the sub-family Termitinae consists of about
270 species. The sub-family Nasutitermitinaeconsistsoffty-sixspecies,which
mainly feed on grass, leaf litter and wood including logs, stumps and standing
dead trees. The genus Trinervitermes specialises in feeding on grass.
The last sub-family is the Macrotermitinae (fungus growing termites)
consisting of over 160 African species, and arguably the most destructive crop/
wood-feeding insects. The main genera include Odontotermes, Macrotermes,
Pseudacantotermes, Microtermes, Ancistrotermes and Allodontotermes.
Odontotermes species consume a variety of plant material and are serious
pests of crops, trees and wood. Some species build massive and tall mounds,
others build low, attened mounds, while others do not build mounds at all.
Macrotermes species dominate the termite fauna in arid environments and build
the most massive mounds that are characteristic of African savannas. While many
Macrotermes species have a narrow range, some such as Macrotermes bellicosus,
Macrotermes falciger, Macrotermes michaelseni and Macrotermes subhyalinus
occur throughout most of the African savanna. Macrotermes bellicosus occurs
from Eritrea in the North to South Africa in the South, from sea-level to
1 800 m,
under moist conditions. Members of the genus Pseudacanthotermes exhibit a
wide range of feeding and nesting habits. The remaining genera (i.e. Microtermes,
Allodontotermes, Ancistrotermes and Synacanthotermes) build subterranean
nests without any surface structure indicating the presence of a colony in the soil.
They feed mainly on wood, litter, dung and occasionally damage trees and crops.
Of these, the most destructive and economically important genus is Microtermes,
which consists of serious pests of trees, wood, crop plants and lawns.
Many species are readily identied using the characteristic of the soldiers’
headsandmandibles. The‘key’usedin the identicationandclassication of
termite species is thus based on morphology and characteristics of the soldier
caste (see Appendix 1).
About 1 000 termite species are known to occur in sub-Saharan Africa and
most of them are of great importance in recycling plant material. A few of them
(about 5 per cent) are serious pests of crops and tree species. In Africa, the
reputation of termites as pests is coupled with the presence of large mounds in
cropeldsorclosetotrees.TermitesarewidespreadinAfrica,restrictedmainly
by desert areas and the lower temperatures found at higher altitudes. There are
approximately 20-50 damaging termite species in savanna and forest ecosystems
in the family Termitidae (fungus-growing termites). The Termitidae contain over
80 per cent of the known genera and over 70 per cent of the known species. In
Zambia, for example, there are about twenty-seven genera with forty-one fully
identiedspecies(Nkunika,1982).Doubtlessly,severalmorespeciesandafew
more genera could be added in future research before the list is complete.

18
Table 1: Approximate Number of Termite Species with Known Distribution in African
Savannas and their Feeding Habits (modied from Sileshi et al., 2010)
Family Sub-family Genus Number
of species
Feeding/damage
Kalotermitidae Kalotermitinae Bicornitermes 4Dry Wood
Biditermes 7Dry Wood
Cryptotermes 4Dry Wood
Epicalotermes 4Dry Wood
Kalotermes 3Dry Wood
Neotermes 17 Dry Wood
Hodotermitidae Hodotermitinae Hodotermes 2 Grass, crops
Termopsidae Termopsinae Microhodotermes
Porotermes
1
3
Grass, litter
Damp wood
Rhinotermitidae Coptotermitinae Coptotermes 6 Wood, trees, crops
Psammotermes 4 Wood, trees, crops, dung
Rhinotermitinae Schedorhinotermes 2 Wood
Apicotermitinae Acholotermes 4 Soil
Termitidae Termitinae Amitermes 17 Soil, wood, litter, dung
Angulitermes 6 Soil, wood
Cubitermes 70 Soil, dung
Microcerotermes 46 Soil, wood, crops, dung
Noditermes 9 Soil
Ovambotermes 1 Wood
Procubitermes 20 Wood, dung
Promirotermes 10 Soil, wood, dung
Termes 7 Soil, wood, dung
Nasutitermitinae Baucaliotermes 2 Grass, litter, wood, dung
Fulleritermes 5 Wood, litter, dung
Nasutitermes 10 Wood, trees, crops
Rhadinotermes 1 Wood, litter
Spatulitermes 1 Wood
Trinervitermes 17 Grass, litter, wood
Macrotermitinae Acanthotermes 1 Wood
Allodontotermes 3 Wood, trees, crops, dung
Ancistrotermes 10 Wood, trees, crops, dung
Macrotermes 14
Wood, grass, trees, crops,
dung
Microtermes 42
Wood, grass, trees, crops,
dung
Odontotermes 78
Wood, grass, trees, crops,
dung
Protermes 5 Wood
Pseudacanthotermes 8
Wood, grass, trees, crops,
dung
Synacanthotermes 3 Wood

19
2.5 Termite Food
The normal food of most termites consists of cellulose (dead wood), fungi
or soil. Some termites feed directly on dead plant tissue; others collect plant
materials on which to grow fungus gardens in their nests and the rest consume
huge amounts of organic matter to derive nutrients. The fungus-growing termites
are the most troublesome in agroforestry. They feed on organic material such as
crop residues, mulches and soil organic matter (humus). However, when this
type of food is not available, they will feed on live plant material including crops
such as groundnuts, millet, maize, cassava and trees. Harvester termites collect
live green plant material and cause damage to pasture grasses, crops and tree
seedlings. They readily attack weak plants that are wilting or damaged. Termite
damageinagroforestryandagriculturaleldsappearstohaveintensieddueto
climate change.

21
Chapter three
BENEFICIAL EFFECTS OF TERMITES
3.1 Livelihood Benets
3.1.1 Termites as Human Food
In many African societies, the winged, reproductive adult termites are regarded
as a delicacy (Sileshi et al., 2009). The queen and soldiers of some species
are occasionally eaten. Local people easily tell the edible termites from those
unsuitable for consumption by vernacular names. The species of termites eaten
in eastern and southern Africa are shown in Table 2 below.
Table 2: Termite Species that are eaten in Eastern and Southern Africa
Species Stage Eaten Country
Hodotermes sp.Nymphs South Africa
Microhodotermes viator Winged adults South Africa
Macrotermes falciger Wingedadultsandqueen Zimbabwe
Macrotermes natalensis Winged adults Zimbabwe
Macrotermes swaziae Winged adults South Africa
Odontotermes badius Winged adults South Africa
Termes capensis Winged adults South Africa
Macrotermes subhyalinus Winged adults and soldiers Uganda
Macrotermes bellicosus Winged adults Uganda
Odontotermes latericius Winged adults Uganda
Pseudacanthotermes spiniger Winged adults Uganda
Odontotermes kibarensis Winged adults Uganda
Pseudacanthotermes militaris Winged adults Uganda
Macrotermes falciger Winged adults Zambia
Termites are commonly dried
and sold, and hence are a
supplementary source of
proteins in human nutrition and
a source of income.
Edible termites on sale at Chipata market (Photo: G.W. Sileshi)

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3.1.2 Termite Mounds as Sources of Mushrooms
There is a mutualistic relationship between some edible mushrooms (genus
Termitomyces) and the fungus growing termites (Macrotermitinae). Hence,
these mushrooms grow mainly around termite mounds. The common mushroom
species include; Termitomyces titanicus, Termitomyces clypeatus, Termitomyces
microcarpus, Termitomyces eurrhizus, Termitomyses letestui, Termitomyces
reticulates and Termitomyces robustus. Termitomyces titanicus is the world’s
largest (up to 90 centimetres in diameter) and one of the tastiest mushrooms.
The indigenous knowledge of mushrooms and their association with termites is
elaborate in many African communities (Sileshi et al., 2009).
A man holding an edible mushroom (Termitomyces)
collected from a termite mound (Photo: Super Stock)
Termitomyces mushrooms on sale at Chipata
market (Photo: G.W. Sileshi)
Small Termitomyces mushrooms around termite
mound (Photo: G.W. Sileshi)
An edible Termitomyces mushroom from
Namibia (Photo: Africa Hunting.com )

23
3.1.3 Termites Improve Agricultural Land
Like earthworms, termites loosen, aerate and enrich the soil by assisting in litter
decomposition and recycling of soil organic matter. They also breakdown and
release organic matter as they digest soil. Recent analyses by Sileshi et al. (2010)
show that mounds of the fungus growing termites have 16 per cent more carbon
content than the adjacent soil. Total nitrogen (N) was also 42 per cent more
in the mound soil compared with the surrounding soil. Termite mounds were
signicantlyenrichedinexchangeablecalcium(232 per cent, more) potassium
(306 per cent more), magnesium (154 per cent more) and sodium (78 per cent
more) than the surrounding soil (Khalil et al., 1990). Termite mound soils were
85 per cent more enriched in cations than the surrounding soil. This is probably
because termites transport cation-rich clay from the subsoil for the construction
of the mounds (Bignell, 2006). Soil pH also increased by 8 per cent in the termite
mounds compared with the adjacent soil (Sileshi et al., 2010). Throughout the
semi-arid regions of Africa, crop growth variability related to termite activity
has been utilised by subsistence farmers in low-risk farming strategies for crop
production. Farmers grow vegetables and cereal crops around mounds to take
advantage of the fertility (Sileshi et al., 2009).
As soil from termite mounds is rich in macro and micro nutrients, African
farmerscrushand use moundsoilasfertiliser in cropelds.Inmany partsof
Africa, farmers apply termite mound soil to the eld where they plant cereal
and legume crops. In Uganda, farmers plant crops such as pumpkins, tomatoes,
onions and maize adjacent to mounds. Similarly, okra, pumpkins, sweet sorghum
andlatecropofmaize,whichrequiregoodwaterandnutrientsupplyaregrown
almost exclusively on termite mounds in Zimbabwe (Sileshi et al., 2009). In
Malawi and Zambia, vegetables are grown on termite mounds while maize is
grown on the soil around them. In southern Zambia and Zimbabwe, farmers apply
termitemoundsoiltotheeld,wheretheyplantmaize,soya beans, cowpeas
and other local cereals and legumes (Siame, 2005; Sileshi et al., 2009). They
do this every three years. It was found that where termite mound soil had been
incorporated, maize harvests were 33 per cent higher than they had been when
inorganic fertilisers were used and the positive effects were long lasting.
3.2 Provision of Ecosystem Services
3.2.1 Termites as Food for Animals
Termites form the major dietary component of many animals. For some animals
such as the aardvark, termites are the main food. Termites are also the most
nutritionally important insects in the diet of chimpanzees and gorillas. During
swarming, termites form a major source of protein rich food for many amphibians,
reptiles, birds and mammals (Sileshi et al., 2009).

24
3.2.2 Termite Mounds as Sources of Minerals
Termite mounds are a source of minerals for animals and humans (Sileshi et al.,
2009). Both domestic and wild animals use mounds as salt-licks because the soil
is rich in essential minerals such as manganese, cobalt, copper and selenium.
Even well fed cattle on commercial ranches cannot resist termite mounds, eating
away the soil to form a grotto and eventually demolishing the whole mound
(Noirot, 1970). Humans also deliberately ingest termite mound soil. This is
calledgeophagy,andithasbeenpractisedbyhumanssinceantiquity.Consuming
soil has often been considered as an aberrant behaviour. However, recent studies
show that this is a perfectly normal behaviour. In Africa, geophagy is common
especially among nutritionally vulnerable populations, pregnant and lactating
women and children. Termitaria are the major sources of the soil consumed by
women and children in Zambia and Zimbabwe (Nchito et al., 2004). There is
also a growing body of evidence suggesting that mound soil can be a source of
irontoanaemic and iron-decientindividuals.Africantermitemounds,which
concentrate minerals up to 500-fold over adjacent soils, probably supply most
or all of the recommended daily allowances of minerals for pregnant women.
Amongthepositivehealthbenets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. Another adaptive
function of consuming termite soil is the ability of clays to adsorb toxins from
plants eaten by humans. This may enable people to rapidly adapt to some of the
many toxic plants (Rowland, 2002).
3.2.3 Termite Mounds are Hotspots of Livestock Feeding
There is growing evidence demonstrating that termite mounds are ‘hotspots’
of foraging by herbivores and ‘keystone’ resources in nutrient-poor rangelands
(Sileshi and Arshad, 2012). For example, in Africa, animals preferentially feed
on termite mounds, compared to the adjacent areas. The higher concentration
of soil nutrients, especially nitrogen, phosphorus and potassium around termite
mounds creates conditions that attract herbivores by providing concentrated
sourcesofhighqualityforage.

25
3.2.4 Termites help Regenerate Degraded Land and Tree Establishment
Termitaria have long been known to have a profound effect on soil, vegetation
composition and structure. In uncultivated land, termite mounds can help in
the establishment of vegetation patches through several mechanisms. Termites
concentrate nitrogen, phosphorus, potassium and bases and elevate clay content
in and around mounds. Increased clay and organic matter around termite mounds
also enhance nutrient and water retention (Sileshi et al., 2010). The mounds
also alter the hydrology and drainage of the site. This can stimulate growth,
diversity and composition of woody plant species (Joseph et al., 2012; Sileshi et
al., 2010). For example, on savanna sites in Burkina Faso, seedling regeneration
was more abundant on Macrotermes mounds than adjacent areas. Acacia spp.
survived better around the mounds than in the inter-mound area at two sites in
Kenya. In the Sahel, manipulation of termite density and activity has been shown
to increase woody plant regeneration and speed the restoration of degraded areas
(Mando et al., 1999). For example, tree species such as Faidherbia albida used in
traditionalagroforestry,deserticationcontrolandre-greeningoftheSahelhave
been shown to regenerate faster around termite mounds (Sileshi et al., 2010).
3.2.5 Termite Mounds are Hotspots of Biodiversity
There are many organisms that live in symbiotic or mutualistic associations with
termites. These include bacteria, protozoa and fungi. Microbial communities
that inhabit termite nests are also of great genetic and functional diversity
(Breznak, 2000). Among the fungi, members of the genus Termitomyces live
in a mutualistic association with the Macrotermitinae, and produce edible
mushrooms. Termite mounds also provide habitat for a variety of termitophilous
animals. These include insects and many other arthropods. Many termite species
that are not mound-builders themselves make their nests within the fabric of
existing termite mounds.
Other animals such as ants, spiders, centipedes and assassin bugs live in or
near the nests and regularly prey on the termites. In addition, birds, reptiles
(e.g., geckoes, lizards and snakes) and mammals preferentially nest in termite
mounds (French and Ahmed, 2010). The Nile monitor buries its eggs in termite
mounds for incubation. Mongooses are largely dependent on termite mounds
for safe den sites.
Plant communities that grow on termite mounds are often different from the
surrounding matrix vegetation (Joseph et al., 2012; Sileshi et al., 2010). Thus
mounds can also be hotspots of plant diversity.

27
Chapter FOur
TERMITES AS PESTS
Termite attacks on annual and perennial crops, especially in the semi-arid and
sub-humidtropicscausesignicantyieldlosses.Itis,however,tobeemphasised
here that not all termites are pests since the normal food of most termites consists
of cellulose (dead wood), fungi or soil. Out of over 1 000 known species of
termites in Africa, only about 5 per cent of these are serious pests of crops and tree
species. Some of the pest species can cause considerable damage in agriculture,
forestryandwoodenttingsin buildings.Harvestertermites collectlivegreen
plant material and cause damage to pasture grasses, crops and tree seedlings.
Hodotermes and Trinervitermes species specialise in grass-eating, which makes
them a serious pest of pasture and rangeland. The fungus-growing termites are
the most troublesome in agriculture and forestry. They feed on dead organic
material such as crop residues, mulches and humus. However, when this type
of food is not available, they will eat live plant material including crops such as
groundnuts, millet, maize, cassava and trees. They readily attack weak plants
that are wilting or damaged.
The greatest pest potential is within the genera Macrotermes, Odontotermes,
Pseudacanthotermes, Ancistrotermes and Microtermes, which consist of serious
pests of trees, structural timber, wood, crop plants and lawns. Macrotermes
species attack plants at the base of the stem, ring-barking or cutting them through
completely. Odontotermes damage is due to feeding either under soil sheeting on
the outer surface of the plants or on the roots.
Microtermes and Ancistrotermes
species attack plants from below
ground by entering the root system
and tunnel into the stem, hollowing it
outandfrequently llingitwith soil.
In Africa, the reputation of termites as
pests is coupled with the presence of
large mounds in crops or close to trees
(Sileshi et al., 2009; 2010).
Termites attack most of the exotic and indigenous tree species used in agroforestry.
Exotic trees such as Acacia and Leucaena species are especially susceptible to
termite damage. Damage to trees and crops is especially severe during the dry
season and dry spells during the rainy season. In eastern Zambia, several farmers
Macrotermes bellicosus damage Eucalyptus
grandis sapling in Uganda (Photo: P. Nyeko)

28
who planted Acacia crasicarpa as woodlots completely lost them due to termite
attack (Sileshi et al., 2008).
In Busia district of Western Kenya,
complete destruction of Casuarina
seedlings were reported. In Rwanda,
it is reported that planting of trees
is badly hampered by termites. In
eastern Uganda, losses of up to 100
per cent have been attributed to
termites on Grevillea robusta (Nyeko
and Olubayo, 2005). In southern
Zambia, termites cause serious
economic damage to agroforestry
plantations. The damage that occurs
in this area ranges from 20-80 per cent
particularly during drought periods.
Fruit tree orchards of citrus, mangoes, avocados, guavas, macadamia and
indigenous fruits, particularly those planted to newly cleared and prepared
lands, are vulnerable to attack by termites. Sometimes their attack results in the
total destruction of entire orchards. The termites implicated are Macrotermes,
Odontotermes and Ancistrotermes species. Trees are most at risk during the
seedling stage and the rst year of establishment in the eld. Seedlings are
either cut just below or above the soil surface. In the latter case, termites gain
access from soil-covered galleries impinging on the base of the plant. Usually,
the seedlings are completely severed, resulting in lowered plant populations.
Damage to mature plants is largely caused by Microtermes and Ancistrotermes
species. These species enter and consume the root system, which directly kills
the plant or indirectly lowers yield through decreased translocation of water and
nutrients. Attack to the root system can also lead to increased susceptibility to
pathogens and other secondary pests.
Termitesalsoinictseveredamagetoagriculturalcropsespeciallygroundnuts,
maize, sugarcane and cassava throughout Africa. Microtermes and Odontotermes
species are the commonest groundnut pests in semi-arid tropical countries of
Africa. The smaller fungus-growing termites (Microtermes spp. in particular),
attack and invade growing groundnut plants through the roots and stem near
ground level, hollowing them out and causing the plants to wilt and die. Stem
bases are sometimes severed by Macrotermes, with 25-100 per cent of plants
beinglostinthisway.Asthecropripens,theouterlayersofthepodsarescaried
by termites, allowing contamination of the seed with soil fungi (e.g., Aspergillus),
which produce aatoxins. Infected plants are not obviously diseased and are
Macrotermes bellicosus damage on Grevillea
robusta sapling in Uganda (Photo: P. Nyeko)

29
frequentlyharvestedwiththefungiandcontaminatetherestofthecrop.InSouth
Africa, up to 30 percentofthe groundnutpods maybescaried.Microtermes
alsopenetratetheshelltofeedonthesoftinnerlining,llingthepodwithsoil.
This form of attack leads to additional loss through premature germination of
kernels.Stacksofplantsleftdryingintheeldsarealsofrequentlyattackedby
Odontotermes, which can cause 30-40 per cent crop loss at this stage. Termite
damage is generally most serious towards the end of the growing season, just
prior to harvesting, with drought being an aggravating factor.
Among cereal crops, maize is most seriously damaged by termites in Africa.
Microtermes and Ancistrotermes attack seedlings and mature maize plants while
Macrotermes species cause damage to seedlings. Odontotermes, Allodontermes
and Pseudacanthotermes can consume the entire seedlings. Maize plants attacked
early in the season can compensate damage with new growth. Lodging of mature
maize plants is one of the obvious symptoms of termite damage. Grain from
lodgedmaizeorgroundnutmaybeinvadedbysoilfungithatproduceaatoxins.
Maize yield losses of between 30-60 per cent have been reported in some parts
of Africa (Sileshi et al., 2005; 2009).
The most damage to sugarcane is done by Amitermes, Pseudacanthotermes,
Macrotermes, Odontotermes, Microtermes and Ancistrotermes. Yield losses of
18 per cent were reported in Sudan, 5-10 per cent in the Central African Republic,
and germination failure of up to 28 per cent was reported in Nigeria. The most
common damage to sugarcane is the destruction of the setts (planting material).
Cassava, which is grown from
stem cuttings is consistently
attacked as seed pieces by
Macrotermes,
and Amitermes,
a species that is predominantly
a root-feeder. Other species like
Ancistrotermes, Macrotermes
Odontotermes, Microtermes
and Pseudacanthotermes also
damage the maturing crops and
trees by hollowing out stems at
ground level. Allodontermes,
Ancistrotermes, Hodotermes,
Microtermes and Odontotermes
often damage cotton, especially
in the drier parts of Africa.
Macrotermes falciger damage on cassava cutting
in Magoye, Zambia (Photo: P.O.Y. Nkunika)

31
Chapter Five
TERMITE MANAGEMENT
We would like to make a distinction between termite control and termite
management. Pest control refers to the elimination of a species perceived as a
pestthroughmeasures such asapplicationofpesticides, re,etc.Termites are
much maligned as pests. Destruction and poisoning of their mounds have been
actively pursued as the main control measures in cropland as well as rangelands
(Sileshi et al., 2009). Unfortunately, control measures are usually taken based on
wrong assumptions as demonstrated by the case study below.
Case Study
Wood (1991) documents a good example of a failed attempt to control termites in
rangelands in eastern Ethiopia. Farmers and government ofcials were reluctant
to accept overgrazing as the primary cause of rangeland denudation. Instead, they
regarded termites as the major problem and launched campaigns to poison mounds
using pesticides. In two districts alone, over 600 000 Macrotermes mounds were
poisoned with 12 000 kg of aldrin at the cost of over 200 000 man-days, which turned
out to be ineffective.
On the other hand, management focuses on long-term prevention or reduction
of pest damage rather than eliminating the pest. Appropriate management of the
cropandpreventionofpestsfrombecomingathreataretherstlinesofaction
inmanagement.Thisisusuallyachieved throughacombination oftechniques
such as habitat manipulation, cultural practices and use of resistant varieties.
Pest control is instituted only when preventive methods are no longer effective
or available. It is futile and counterproductive to attempt to control termites in
agroforestry. But it is possible to manage them.
It is important to note that there is no single method that can be universally
recommended for termite control. Methods of control are more effective if used
in conjunction with each other, with maximum use of local knowledge and
resources. In that sense, we advocate termite management rather than controlling
them.Therstpartofthischapter,therefore,focussesongeneralprinciplesthat
help in reducing termite damage. The recommendations given are not restricted to
aspecicspecies.Theyaremeanttoreducetheimpactofpestsandotherfactors
on the growth of the plant. The second part focuses on specic management
methods tested on some termites and found effective.

32
5.1 General Principles for Reducing Termite Damage
The suggestions given below are based on review of a wide range of literature.
Some of them come from individual research studies and have not been widely
tested. Most of the principles may not necessarily reduce termite numbers, but
only reduce damage to trees and crops.
Good Nursery Management
Seedlingqualityrepresentsthe most importanteconomicaspectof forestation
andagroforestry.Tomeetfuturedemandsforplantingmaterial,highqualityand
quantityoftreeseedlingsmustcontinuetobeavailabletothefarmers.Increased
planting of agroforestry species confronts nursery managers with a wider array
of potential pest problems. When major problems do occur, nursery owners
can utilise integrated pest management practices. This means the integration of
suitable techniques and procedures into one concerted, harmonious effort for
effectiveandefcientcontrolofnurserypests.IntegratedNurseryManagement
(INM) is the most practical and ecologically sound approach for control of
nursery pests because it also involves all other nursery management procedures.
INM is accomplished through good nursery management (Jaenicke, 1999) in
combination with tactics that prevent pest damage from reaching economically
damaging proportions, while protecting the environment from potential hazards.
These tactics include preventing pests (exclusion) from multiplying in the
nursery, use of good and healthy tree seeds, planting seeds at the right time and
density, watering, sanitation, and controlling pests through chemical or biological
means. Hence, INM is not only aimed at termites but at all other problems
(including pests) that induce stress, leading to termite damage once seedlings are
out-planted.TheobjectiveofINMistoproducegood-qualityhealthyseedlings
consistentlyandprotably.
Protecting Tree Seedlings
Therstyearisacriticalperiodwhentreeseedlingsaremostatrisk.Ingeneral,
termite attack becomes minimal after canopy closure. Therefore, management
may be mandatory during the nursery and seedling stages, with some exceptions
in a few older plants. The presence of termite mounds or foraging activities in
theeldin well-establishedplantationsdoes notnecessarilyindicatea termite
hazard.

33
Providing Substitute Sources of Food
There is a common belief that dead plant materials attract termites to live trees.
This is a myth because fungus-growing termites, which are the most serious
pests of trees, prefer to eat dead plant material. The solution is rather to provide
termites with an alternative source of food such as mulching with wood and dry
leaves. This may not only divert termites from live plants but also add organic
matter to the soil. Therefore, farmers should avoid leaving bare, dry soil around
seedlings or trees. It is important to leave as much plant debris (dry wood, leaves,
grass) as possible on the soil surface when preparing planting sites. Mulching
with items such as hay, manure, wood shavings, wood ash or maize stover has
been shown to dramatically decrease termite attack on live plants. Termites are
attracted to the mulch rather than the crop or tree. Vetiver grass leaf mulch has
been shown to prevent termite attack around the base of trees.
Maintaining Soil Organic Matter
The damage by many termites becomes severe on soils with low organic matter
content. This is because such soils do not contain enough food for termites to
live on and they resort to feeding on living plant material. Adding compost or
well-decomposed manure to the soil and sowing green manure legumes and
cover crops helps to increase the organic matter in the soil. However, attack
may increase if there is an abundance of undecomposed organic matter such as
manure that attracts root feeding termites such as Odontotermes. This may also
increase nesting by ants, which are the worst enemies of termites.
Ensuring Overall Plant Health
Termites rarely attack healthy plants but may do so following weakening of
plants. Anything that weakens the plant such as drought, neglect of cultural
practices, low soil fertility and lack of essential nutrients, overcrowding and
competition, weed infestation, injury by re and insects or fungal attack will
predispose the plant to termite damage. In dry areas, seeds should be sown at the
beginning of the wet season to give the plants a chance to establish themselves
andremainhealthyinthe eld. It is also recommended thatonlyhealthyand
vigorouslygrowingtreeseedlingsbetransplantedintotheeld.Plantswhichare
suffering from disease or lack of water are generally more susceptible to termites
than healthy plants. It is, therefore, important that plants are kept healthy and not
stressed. The following practices are recommended to reduce stress:

34
Watering
Give nursery stock enough water just before planting out. Plants become
susceptible to termite attack during the dry season if they are allowed to dry due
toinadequateorincorrectwatering.Waterseedlingstothepointofwaterlogging
withtherstsightoftermitedamage.
Root Pruning
Root pruning is a necessary operation in tree nurseries. After root pruning, allow
enough time for recovery of damaged roots before transplanting seedlings. In
nurseries,itisimportanttoschedulerootpruningtoallowsufcientrecoveryand
repair of damaged tissues before transplanting. Field root pruning may cause less
stress to trees if conducted during rainy periods. Alternatively, raising seedlings
on a raised bed ensures air-pruning, hence reducing the risk of termite damage.
Transplanting
Transplant seedlings on time (at the beginning of the wet season) and when the
soilissufcientlywetorwhenthegroundismoist(ininstanceswhereirrigation
systems are practised).
Observing Planting Date
In unimodal rainfall and dry areas, it is recommended that seeds should be sown
at the right time to give the plants a chance to establish themselves before the
onset of the dry season. As a general rule, begin planting out seedlings with the
rstgoodrainsorwhenthesoiliswettoadepthof20-30cm.
Correct Planting Density
Plant tree seedlings using the recommended densities to reduce competition.
Timely Weeding
Weedtheeldpromptlytoreduceweedcompetition.
Avoiding Planting Susceptible Species
In areas where termite damage is severe, it is prudent to opt for species that are
tolerant or resistant to termites and avoid those that are prone to termite damage.
However, resistance is a relative concept, which depends on the tree species,
origin of the tree, age and condition of the tree, termite species and soil and
climatic conditions where the tree is growing. Some indigenous tree species are
more resistant to termite attack than exotic species. There is little knowledge

35
about crop resistance to termite attack. However, in general, indigenous crops and
tree species are more resistant to termites than exotic species because indigenous
species are better adapted to the local ecology. For instance, in Africa, sorghum
and millet are more resistant to termites than maize and cowpea. Bambara nuts
are not attacked while groundnuts suffer serious damage.
Encouraging Natural Control of Termites
Termites have a wide variety of enemies including black ants, beetles, bugs and
spiders. Ants are the greatest enemies of termites in all regions of the world.
Decomposing materials such as dead animals, sh bones, manure or sugary
substances (e.g., sugarcane husks) attract ants. This has been traditionally applied
by farmers in Africa (Sekamatte, 2000; Sileshi, 2008). A recent test in maize
eldsinUganda hasshownthatprotein-based baitssuchas molassesandsh
bonesattractsignicantnumbersofantsandmoreantsestablishnestsnearmaize
plants (Sekamatte, 2000). This in turn reduced termite damage and increased
grain yields. Other predators of termites include frogs, lizards, snakes, birds
and bigger animals such as aardvarks, pangolins, bats, monkeys and humans.
Encouraging this kind of wildlife will help to reduce the number of termites.
Bushes and trees are a home for many of these useful creatures. Areas of natural
habitatshouldalwaysbeleftaroundeldswherecropsandtreesaregrown.If
these areas are destroyed, then there is an imbalance between the populations
of predators and termites. Agroforestry can also increase natural enemies of
termites and reduce termite damage on crops.
Intercropping and Mixed Cropping
These are the most effective cultural practices used by small-scale farmers in
sub-SaharanAfricatomanageinsects thathave specichost ranges.However,
controversial results have been reported for termites. For example, intercropping
maizeandbeans resultedinsignicant reductionoftunnelling bytermitesbut
did not reduce termite damage on the plants. On the other hand, intercropping
in forestry has been suggested as a means of retaining termite diversity in the
crop in order to prevent them from achieving pest status. Certain grasses are
intercropped with different crops in West Africa to repel termites. Mixing trees
with crops also can reduce termite damage to either the crop or tree component.
For example, in eastern Zambia, planting maize between rows of tree stumps
reduced termite damage on maize (Sileshi et al., 2005).

36
Crop Rotation
Planting the same crop on the same land year after year reduces soil fertility and
structure. Crops growing in such conditions will be weaker and susceptible to
termites. Crop rotation means that different crops are grown on the same piece
of land each year. This can prevent pest and disease build up and also help the
soil to recover nutrients.
5.2 Specic Control Methods
5.2.1 Chemical Control
In the past, chemical control of termites was largely based on the use of
organochlorine insecticides such as lindane, aldrin, dieldrin, chlordane and
heptachlor. These were applied as seed dressing or to the soil in planting holes or
furrow treatments or poisoning the mound. However, termites soon developed
resistance to these chemicals. Resistance to aldrin, for instance, had been
reported as early as the 1970s in Eucalyptus plantations in Zambia. Following
increasing restrictions on the use of persistent organic pollutants, less persistent
insecticides such as the organophosphates (Chlorpyrifos, Isofenphos), carbamates
(Carbosulfan, Carbofuran), and pyrethroids (Permethrin, Decamethrin) have
been used as alternatives; but their low persistence often necessitates repeated
applications. Recently, controlled-release formulations of some non-persistent
insecticides were tried and found to be effective and long lasting. However, these
formulations are not cost-effective for the majority of low-income farmers in
developing countries.
5.2.1.1 Alternative Control Options
There are a number of alternatives to using chemical pesticides for termite control.
Some of these methods work within the natural system and help promote natural
pest control mechanisms. For instance, organic control methods do not pollute
theenvironmentandarenotharmfultobenecialanimals,ortothepeopleusing
them. Organic methods aim to use locally available materials and do not rely on
importing expensive materials from elsewhere. Organic methods are thus cheap
and easy to use. These methods regulate termite numbers rather than eliminate
themsothatthebenetsprovidedbytermitesarenotlost.

37
Plant Preparations (Botanicals)
The pesticidal properties of many plants have been known for a long time
and natural pesticides based on plant extracts such as rotenone, nicotine and
pyrethrum have been commonly used in pest control (Stoll, 1986). There are
many other plants with insecticidal and repellent properties. However, the
effectiveness of plant extracts vary depending on the target insect, concentration
used, the plant part, the maturity of the plant and the condition under which it
has been grown. Most farmers like to see pests drop dead right away. However,
plant extracts including those of the famous neem do not have this effect. Their
main effect is as a repellent or antifeedant, which means insects avoid the treated
plants. Different plant parts may be used as a natural insecticide. Concoctions of
plant parts, such as toxic fruit juices, pulps or shavings can be applied directly.
The advantage of plant extracts is that, unlike synthetic pesticides, they leave no
residue on crops as they breakdown within a few days after application.
Farmers have traditionally used plant extracts for millennia and a rich
indigenous knowledge still exists in some communities. Farmers in different
areas use different methods of preparing plant extracts. Here, we will only
provide a few examples of plants commonly used by farmers to control termites
(Sileshi et al., 2009).
Tephrosia vogelii
Leaves and pods are crushed into a ne powder and mixed with water, then
sprayed on affected plants.
Lantana camara
Leaves are boiled in water for 30 minutes, and then sprayed on the crop.
Neem (Azadirachta indica)
Leaves or fruits are crushed and soaked in water; the solution is sprinkled on the
crop.
Euphorbia tirucalli
Roots are soaked in water for 24 hours; the solution is sprinkled on insects and
on the crop.
Bobgunnia (Swartzia) madagascariensis
Pods are crushed and put in water, then sprinkled on the crop.

38
Melia azedarach
Roots and bark are soaked in water for 24 hours and then sprayed preventively
or directly on attacking termites.
The list of plants above is not exhaustive. There are probably many more plants
that could be locally available and as effective as the ones above. Appendix 2
gives examples of plant species used in Zambia, along with brief descriptions
of the method of extraction and application. These are farmers’ recipes and may
not be effective under all conditions. The limitation of farmers’ recipes is that
different farmers give different versions. Therefore, users should carry out their
own experiments to nd the plants and application rates most appropriate to
theireldconditions.Mostplantmaterialsalso breakdown rapidly in the soil
and do not give prolonged protection from termite attack. Unlike conventional
pesticides, little is known about the active ingredients of local indigenous
pesticides and their mode of action (see Appendix 3). In addition, the hazards
they present to humans and the environment is often unknown. Therefore, greater
careisrequiredintheiruse.
5.2.1.2 Microbial Preparations
Preparations of the fungus Metarhizium anisopliae has recently been developed
for control of termites in buildings in the USA, Brazil and Australia. Colonies of
some species could be killed when nests were inundated with conidia. Research
conducted at the International Centre of Insect Physiology and Ecology (ICIPE)
has indicated the effectiveness of the fungus for the control of termites in pasture,
nursery trees and mounds in Kenya (ICIPE, 1997). The potential for control of
termites in maize has been demonstrated (Maniania et al., 2002). The Centre
for Agriculture and Biosciences International (CABI) has a large collection
of Metarhizium anisopliae and Beuvaria bassiana isolates which have shown
varyinglevelsofefcacyagainsttermites.CABIandICIPEhavealsodeveloped
formulations, storage and application techniques for these isolates, which are
effective for protecting tree seedlings especially if applied to potted seedlings.
The fungus formulated as granules and applied as seed treatment reduced plant
lodging and increased yield of maize. Conidia of Metarhizium could be spread
through the colony and nest by contact and grooming between contaminated
and uncontaminated termites. Application methods, including inundating termite
nests or sites of high termite activity with formulated products and use of low
doses of the insecticide imidacloprid in combination with fungus have improved
control of termites.

39
5.2.1.3 Wood Ash
Wood ash is widely used by Zambian small-scale farmers. It may be as effective
as chemical pesticides, but has great advantage in that it is harmless to human
healthandcheaplyobtainedfromrewoodandcropresidues.Abitofsoilis
removed around the plant and then ash is sprinkled around the affected plant.
Ash is alkaline and it may, therefore, repel termites. Besides, wood ash blocks
the tracheae of small insects such as termites, thereby suffocating them.
5.2.1.4 Cow Dung and Urine
Cow dung and urine have been used for termite control by farmers in many
parts of Africa (Sileshi et al., 2009; 2008). In Machakos district of Kenya,
farmers smear cow dung on posts to protect them from termite attack. In Monze
district of Zambia, farmers used fresh cow dung to reduce termite damage to
maize. Similarly, farmers in south-western Nigeria believe that goat and cow
dung reduce termite damage. Reduction in termite damage to rangeland using
cow dung has been demonstrated in an experiment conducted in the ‘Cattle
Corridor’ of Uganda. Cow dung is suspected to reduce termite damage through
the accumulation of organic matter in the soil. However, the precise mechanisms
are not yet known.
5.2.1.5 Destruction of Mounds and Colonies
There are different ways of destroying a colony, including digging the nest
andremovingthequeen,burningwood,grassorcowdungtokill the colony
withsmoke,pouringhotwaterandoodingthenestwithrainwater.Although
destruction of a colony has been advocated by researchers, success has been limited
duetovariousconstraintsincludinglabourrequirementsandlackofknowledge
about termite biology (Sileshi et al., 2009). As this practice is directed towards
mature colonies of the mound-building species, it overlooks species that do not
build mounds (e.g., many Odontotermes and Microtermes spp.). It must be noted
that non-mound building species are often the more serious pests especially in
arid areas. For example, the non-mound building termites such as Microtermes,
Trinervitermes and Microhodotermes spp. destroy rangeland much more than
the mound building termites, including Macrotermes and Odontotermess pp.
Thosethatbuild mounds arealsosubterraneanfor the rstfewyears.Even if
mature colonies are killed, the immature colonies could spread to take over the
area. Many farmers also do not destroy the mounds even when they are live or
veryeasytoatten.Somefarmersalsoresistdestructionofmoundsastheyare
considered sacred places among many communities in Africa.

41
Chapter Six
CONCLUSION
The key message is that not all termites are pests. It must be noted that the pest
activityisapartandparcelofthetermite’sbenecialroletovariousecosystems.
Therefore,itisimportanttoassessthecostsandbenetsoftermitesbeforeany
control measure is taken against the insect. Since not all termite species are
pestiferous, control measures that target only damaging species could provide
the best option. Local communities have comprehensive indigenous knowledge
of termite taxonomy, ecology and apply various indigenous control practices.
These must be strengthened in the face of global climate change and the world
ban on persistent organic pollutants.
We would like to emphasise that it is futile and counterproductive to attempt
to control termites in agroforestry. But it is possible to manage them through a
combinationoftechniquesasoutlinedinthisbook.Chemicalcontrolshouldbe
instituted only when preventive methods are no longer effective or available.
Thisbookraisesmorequestionsthananswers.Thiscallsforfurtherresearchin
this much neglected, yet a very important area in tropical agroforestry.

42
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45
APPENDICES
Appendix 1
Key to the termite genera based on the soldier caste (After Mitchell, 1980)
1. Pigmented compound eyes present.......................................................
Pigmented compound eyes absent.........................................................
2
3
2 Compound eyes conspicuous. Head rounded. Pronotum
with broad median anterior lobe.............................................................
Compound eyes small. Head rectangular. Pronotum without median
anterior lobe...........................................................................................
Hodotermes
Epicalotermes
3Head drawn out into long conical tube. Mandibles reduced........................
Head not drawn out into long conical tube. Mandibles well de velop ed…..
30
4
4 Fontanelle absent…………………………………………...................
Fontanelle present……………………………………….....................
5
7
5Headsquare,frontofheadvertical.Mandiblesshortandangledinthe
middle ………………………………………........................................
Head rectangular, front of head not vertical, mandibles well developed.
Pale, unpigmented eye spots present ………….....................................
Cryptotermes
6
6Pronotum large and kidney shaped with sides evenly rounded into poster ior
margin………………………………….................................................
Pronotum smaller with sides straight and almost parallel….....................
Neotermes
Biditermes
7Pronotumatwithoutanteriorrobes………....................……………..
Pronotumneverat,oftensaddleshaped……………….......................
8
10
8Head pear shaped. Fontanelle very well developed, directed forwards.
Mandibles without marginal teeth. Soldiers monomorphic……………
Head not pear shaped. Fontanelle on top of head. Mandibles with
marginal teeth. Soldiers dimorphic or polymorphic...............................
Coptotermes
9

46
9Headlongwithparallelsides.Leftmandibleswithveormoremarginal
teeth, right mandible with three marginal teeth grouped about half way
along. Soldiers polymorphic....................................................................
Head round. Left mandible with two well developed. Marginal teeth,
right mandible with only one marginal tooth. Soldiers dimorphic……
Psammotermes
Schedorhinotermes
10 Labrum more or less tongue-shaped, rounded or pointed, with or
without hyaline tip………………………………….............................
Labrum of various shapes, square, emarginated or trilobed, without
hyaline tip…………………………………………..............................
11
18
11 Mandibles with conspicuous marginal teeth…………….................…
Mandibles without marginal teeth or with microscopic serrations
only ........................................................................................................
12
14
12 Single large marginal tooth in right jaw. Several smaller marginal teeth
in left jaw…………………………………............................................
13
13 Mandibles sickle shaped. Jaws symmetrical with single marginal tooth
in each……………………………………… ........................................
Mandibles not sickle shaped with or without marginal tooth in right jaw.
Dentition asymmetrical……………………….......................................
Amitermes
Odontotermes
14 Two spine-like points projecting forwards from front edge of Pronotum.
Soldiers dimorphic…………...................................................................
Pronotum without spine like projections. Soldiers monmorphic or
dimorphic………....................................................……………………
Pseudacanthotermes
15
15 Labrum without hyaline tip. Large species. Soldiers dimorphic……....
Labrum without hyaline tip. Small species. Soldiers monomorphic or
dimorphic…….………………...........................................................…
Macrotermes
16
16 Head and long rectangular. Mandibles thick with minute serrations
along inner Margin……………….........................................................
Head rounded. Mandibles slender without serrations along inner
margin ....................................................................................................
Microcerotermes
17
17 Soldiers dimorphic. Mandibles of major soldiers strongly curved
inwards at tip………………………......................................................
Soldiers monomorphic. Mandibles curved in lightly at tip ..…..…......
Ancistrotermes
Microtermes

47
18 Contents of the abdomen light grey to almost black..............................
Contents of the abdomen white to creamy yellow..................................
19
24
19 Labrum abnormally large, emarginate, bilobed with thick rounded
lobes .......................................................................................................
Labrum normal size…………………………………............................
Euchilotermes
20
20 Mandibles massive and powerful………….......................…………..
Mandibles not abnormally heavy……………………..........................
21
22
21 Mandibles short and thick. Labrum triangular, about as long as broad,
tapering to rounded acute angled pointed…………………………….
Mandibles long and thick with sides evenly and convexly curved from base
totip,thecuttingmarginsnelyserrated……………….......…………
Apicotermes
Ovambotermes
22 Labrum forked, deeply bilobed………………………..........................
Labrum not deeply bilobed…………………………….........................
Cubitermes
23
23 Mandibles straight, shorter than length of the head, curved inwards at
the tip. Labrum as broad as long, very slightly emarginated, the corners
drawnoutintwonepoints……………………………………….......
Mandibles long and slender with the inside of the tips attened and
adapted for ‘snapping’. Labrum longer than broad with concave tip....
Crenetermes
Termes
24 Mandibles grotesquely asymmetrical, the left with a cork screw-like
twist, the right straight….......................................................................
Mandibles symmetrical…………………………….............................
Pericapritermes
25
25 Gulawithdeniteridgeorswelling…………………...........................
Gula without ridge or swelling………………………..........................
26
27
26 Eminence on gula a rounded knob. Labrum deeply forked …………
Eminence on gula with two spines directed forwards and outwards.
Labrum wider than long, anterior margin slightly concave with the
corners drawn out to points…………………………………………...
Noditermes
Unguitermes

48
27 Mandibles, in plan view, more or less straight but slightly angled
outwards, the inner faces of the tips slightly cupped and adapted for
‘snapping’ ............................................................................................
Mandibles, in plan view, curved inwards and adapted for biting …….
28
29
28 Inside view the front of the head drawn out into a protuberance ending
in an acute angled point. Sides of head more or less parallel. Mandibles
about as long as the head including the frontal projection……............
Protuberance on front of head in lateral view usually rounded but may
havearatherattenedfacet.Sidesofheadusuallyconvexbutmaybe
attened.Mandiblesusuallydenitelylongerthanhead……………...
Angulitermes
Promirotermes
29 Front of labrum almost straight, slightly emarginated.
Mandiblesverynelystriated………………………….....................
Front of labrum slightly trilobed.
Mandibles not striated………………………………………………
Lepidotermes
Basidentitermes
30 Head capsule, in plan view, triangular, tapering smooth
from near the back to tip of ‘nose’…………………….....................
Head capsule, in plan view, not triangular, the ‘nose’ part being
distinctly more narrow than the head .......................................…….
Mimeutermes
31
31 Head constricted behind antennae…………………….......................
Head not constricted behind antennae ......……………......................
32
34
32 Head and ‘nose’ covered with minute spatulate bristles ……………
Bristles on head normal, with pointed tips……………......................
Spatulitermes
33
33 Colour of head capsule orange or reddish, darker on ‘nose’………….
Colour of head capsule septa brown. ‘Nose’ darker’ with reddish tip ...…
Fullerritermes
Rhadinotermes
34 Soldiers monomorphic………………………………….
Soldiers dimorphic……………………………………..
Nasutitermes
Trinervitermes

49
Appendix 2
Plant species used in Southern and Central provinces of Zambia (Nkunika,
1998; Nkunika, 2002)
a. Muyongolo (Bobgunnia madascariensis)
Grind ten dry pods in a mortar and mix into 2.5 litres (Kachingubuli) of
water. Leave the mixture for 24 hours. Shake the mixture well before
using. Some foam should be seen on top of this mixture. Pour or splash
the mixture directly on the bases of the affected tree. Do this for 5 days,
after that continue with normal watering. This concoction was very
effective against termites.
b. Muleyembezo (Crossopterix febriguga)
Dig and cut 10 pieces of roots about (15 cm) long. Peel off the bark
and grind. Using two handful of the crushed powder soak together with
roots in 5 litres (Chingubuli) of water. Leave for 24 hours. Pour the
mixture around the base of the tree. This mixture is very effective against
termites.
c. Mumba (Jurbanadia globiora)
Collect and put a handful of leaves in 2.5 litres of water and soak them
for 24 hours. Sieve the mixture and pour around the bases of the effected
tree. The mixture was effective against termite pests.
d. Mululwe (Cassia abbreviata)
Get two hands full of the bark of mululwe, grind and mix with 5 litres
of water together with the dry preparation of muyongolo. Pour on the
affected tree. This concoction is very effective against all termite pests.
Mululwecanalsobeusedonitsownandgetequallyeffectiveresults.
e. Ntuntulwa (Solanum incanum)
 Cut the fruit of Ntuntulwa into quarters or halves and pound. Put the
pounded fruit into a container and add 2.5 litres of water. Leave the
mixture to stand for 12 -24 hours. Shake the mixture before use. Pour or
splash the mixture on the affected plant. This was reported to be effective
against termites.
f. Kanunka (Bidens pilosa)
First collect a cupful of the mature seeds. Mix with water, boil for ten
minutes or soak for 24 hours. If boiled, allow to cool, and then add 1 litre
of water and a teaspoon full of soap. Spray immediately. This mixture
was less effective against termite pests.

50
g. Mbono (Ricinus communis)
Soak green seeds and leaves in water for 24 hours. Filter and spray
or alternatively put green seeds into hole around the target plant. The
concoction deters the termites.
h. Mbala (Euphobia tirucalii)
Pound the leaves; soak in water for 24 hours and spray. The mixture
should be slightly milky before spraying. Though effective against
termites, this plant is toxic to humans and must be treated with caution.
i. Ububa (Tephrosia vogelii)
Collect 1kg of fresh leaves and grind and soak in 20 litres of water.
Leave for 24 hours. Strain and then spray on the affected plant. Though
effective against termites, this is a recent introduction and most farmers
are not familiar with it.
j. Impili- pili (Capsicum frutescens)
Grind 500g of chilly fruits and soak in 20 litres of water for 24 hours.
Sieve the mixture and use as a spray. This concoction was very effective
on tree seedlings.
k. Muwi (Strychnos cocculoides)
Remove bark and pound together with fruits. Soak in water for 24 hours.
Sieve the mixture. Measure 500 ml and dilute to 10 litres of water. Pour
solution around stem base of affected plants.
l. Mulombe, Muzwamalowa (Xerodenis stuhmaanii)
Strip the bark and soak overnight. Pour solution over affected area. This
mixture was effective depending on the preparation.
m. Tombwe (Nicotiana tabacum)
Crush 3 kg fresh leaves, soak in 20 litres of water and add soap. Use the
mixture to spray around the plant or crush dry leaves of tobacco and add
fresh chillies and soak in water. Use mixture to sprinkle affected plant.
The termites were killed instantly.
n. Muntamba (Strychnos spinosa)
Pound fresh fruits and soak in water and leave overnight. Spray to the
affected area.
o. Sozwe (Cubonia glauca)
Spread leaves or roots over and under maize. Highly poisonous and
repellent to termites. It should not be handled by children or animals.

51
p. Paw Paw (Carica papaya)
 Add 1 kg of nely shredded leaves to one litre of water and shake
vigorouslythenlterandaddfourlitresofwater,twospoonsofparafn
and 20 g of soap. Spray around the bases of the tree. This concoction has
a repellent effect against termites.

52
Appendix 3
Active principle and mode of action of some indigenous plants with pesticidal
properties found in Zambia
Vernacular
Name
Botanical Name Active Principle Mode of Action
Muyongola Bobgunnia
(Swartzia)
madascariensis
Plant contains
saponins
Pesticidal, Repellent
Mbala Euphoria tirucalii Nicotine, Tar Insecticidal, toxicant against
termites
Tombwe Nicotiana tabaccum Notidentied Repellent to termites
Ntuntulwa Solanum incunum Ricin Toxicant to termites
Mbono Ricinus communis Ricin Toxicant to termites
Mutandezyelo Targetes minuta Terpine Insecticidal, Repellent to
termites