5Weeds, Diseases, Insect Pests, and Tri-Trophic Interactions in Tropical Agroforestry

22 Pages·2007·0.2 MB·English

Batishetal./EcologicalBasisofAgroforestry 43277_C005 PageProof page 73 25.6.2007 6:29pm CompositorName:BMani
5 Weeds, Diseases, Insect Pests,
and Tri-Trophic Interactions
in Tropical Agroforestry
G. Sileshi, Götz Schroth, Meka R. Rao, and H. Girma
CONTENTS
5.1 Introduction............................................................................................................................73
5.2 Partitioning the Complexity of Pest Interactions...................................................................75
5.2.1 Interactions Betweenthe Plant Community, Herbivores, and their
Natural Enemies.........................................................................................................75
5.2.2 Interactions BetweenHerbivores and Plant Pathogens.............................................78
5.2.3 Interactions Among Herbivores.................................................................................78
5.3 Interactions inSelected Agroforestry Practices.....................................................................79
5.3.1 Sequential Agroforestry Practices..............................................................................79
5.3.1.1 Rotational Woodlots andImproved Fallows..............................................79
5.3.2 SimultaneousAgroforestry Practices.........................................................................81
5.3.2.1 Treeson Cropland......................................................................................81
5.3.2.2 Mixed Intercropping...................................................................................81
5.3.2.3 AlleyCropping...........................................................................................82
5.3.2.4 Multistrata Agroforestry Systems...............................................................84
5.4 Ecological HypothesesRegarding Interactions.....................................................................85
5.4.1 Plant Stress Hypothesis..............................................................................................86
5.4.2 Plant Vigor Hypothesis..............................................................................................86
5.4.3 Carbon–NutrientBalance Hypothesis.......................................................................87
5.4.4 Natural EnemiesHypothesis......................................................................................87
5.4.5 Resource Concentration Hypothesis..........................................................................87
5.4.6 MicroclimateHypothesis...........................................................................................88
5.5 Summaryand Conclusions....................................................................................................89
Acknowledgments...........................................................................................................................90
References.......................................................................................................................................90
5.1 INTRODUCTION
Under the International Plant Protection Convention, a pest is defined as any species, strain, or
biotypeofplant,animal,orpathogenicagentinjurioustoplantsorplantproducts(ISPM,2006).The
coverageofthisdefinitionincludesweedsandotherspeciesthathaveindirecteffectsonplants.This
definitionalsoappliestotheprotectionofwildflorathatcontributetotheconservationofbiological
diversity. Unless otherwise stated, throughout this chapter the term ‘‘pest’’ refers to weedy plants
andparasitichigherplants,plantpathogenicorganisms(viruses,bacteria,mycoplasma,fungi),plant
parasiticorpathogenicnematodes,arthropods(herbivorousmitesandinsects),andvertebratepests
(herbivorous birds and mammals)that affect trees and associated crops in agroforestry.
73
Batishetal./EcologicalBasisofAgroforestry 43277_C005 PageProof page 74 25.6.2007 6:29pm CompositorName:BMani
74 EcologicalBasisofAgroforestry
Weeds may be classified as ruderals (annual or biennial plants that primarily infest waste
places),argestals(annualorbiennialweedsofcultivatedlands),andenvironmentalweeds(invasive
alien species). Weeds compete with trees and crops for water, light, and nutrients. Many weed
species also serve as alternative hosts of plant pathogenic organisms and nematodes. Exotic tree
speciesusedinagroforestrycanalsobecomeinvasiveandaffectecosystemfunctionsandbiodiver-
sity. According to a recent estimate (Richardson, 1998), out of over 2000 species used in
agroforestry, some 25 species (1%) are invasive. These include Acacia (8 spp.), Prosopis
(3 spp.), Casuarina (2 spp.), Leucaena leucocephala, Sesbania bispinosa. It must be noted here
thatnotallalienspeciesareinvasive,andnotallinvasivespeciesmaybeeconomicallyimportant.
Transformer species—a subset of invasive plants that change the character, condition, form, or
natureofanaturalecosystemoverasubstantialarea—haveprofoundeffectsonecosystemfunctions
and biodiversity and are invasive (Richardson,1998).
Adiseasecanbedefinedasanyphysiologicaldisturbanceofthenormalfunctioningofaplant
asaresultofadetrimentalinteractionbetweenthepathogen,theenvironment,andthehost(Agrios,
1988).Diseasesaffecttheproductionandutilizationoftreesandcropsbyreducingthehealthofthe
plant and directly reducing yield, quality, or storage life. Plant parasitic nematodes mostly affect
plants by inhibiting root growth, and hence overall plant development, and this usually results in
poorcropperformanceorcompletefailure.Manyplantparasiticnematodesalsointeractwithother
microorganisms such as viruses, bacteria, and fungi in the development of disease complexes
(Kleynhans et al., 1996). Herbivorous mites and insects can physically feed on various parts of
the tree, crop, orboth, and also transmitdiseases.
Inthetropics,weeds,diseases,andinsectpestsareestimatedtoaccountfor13%,13%,and20%
oflosses,respectively(Oerkeetal.,1994).Weedcontroltakesover50%ofthetotallaborneededto
produce a crop. Pests have been cited as one of the factors diminishing the benefits from tropical
agroforestry (Mchowa and Ngugi, 1994; Karachi, 1995; Rao et al., 2000). Unless the biological
constraintsimposedbypestsareremoved,thepotentialbenefitsofagroforestryintermsofincreased
capture and efficient use of resources cannot be translated into economic benefits (Ong and Rao,
2001). If the current enthusiasm of farmers for testing and eventually adopting the various
agroforestry practices is to be sustained, it is essential to know how this practice affects pest
populations and their naturalenemies.
Although the relevance of pest interactions with agroforestry practices has been recognized
manyyearsago(HuxleyandGreenland,1989),veryfewdetailedstudiesoftheirinfluenceontree–
cropinteractionsexist.Thereseemstobemorefocusonpopulationecologyofselectedpestspecies
attheexpenseofecosystemecology.Infact,thereexistcertaingeneralmisconceptions,whichhold
that trees have no or fewer pests and that diversity based on trees reduces pest problems in
agroforestry (Desaeger etal., 2004). Thishas hindered progressin theunderstanding of tri-trophic
interactions in agroforestry. Even in the more recent books on agroforestry (Schroth and Sinclair,
2003;Nairetal.,2004;vanNoordwijketal.,2004),thereislittle,ifany,mentionoftheeffectsof
tree–crop interactions on pests and their natural enemies. In the recent reviews, Day and Murphy
(1998) and Rao et al. (2000) dealt mainly with insect pests affecting agroforestry trees and their
management. Schroth and coworkers (2000) dealt with insect pests and diseases in agroforestry
systems of the humid tropics. The review by Gallagher et al. (1999) and Ong and Rao (2001)
focused on managing tree–crop interactions in relation to weeds. Desaeger et al. (2004) dealt with
nematodesandothersoil-bornepathogens.Thereviewontheeffectoftreesonabundanceofnatural
enemies(Dix et al.,1995) focusedon agroforestry systems ofthe temperate zone.
Though complex interactions are known to occur between various categories of pests (e.g.,
weeds, pathogens, nematodes, insects, etc.), the nature of such interactions are poorly understood
and little quantified in tropical agroforestry (Hitimana and McKinlay, 1998). This work is the
first attempt to draw together information on the different categories of pests and natural enemies,
and apply the knowledge to the challenges of pest management in tropical agroforestry. In this
chapter, an extensive review of literature pertinent to tree–crop interactions and pest risks in
Batishetal./EcologicalBasisofAgroforestry 43277_C005 PageProof page 75 25.6.2007 6:29pm CompositorName:BMani
Weeds,Diseases,InsectPests,andTri-TrophicInteractionsinTropicalAgroforestry 75
agroforestrywasconducted.Inviewofthevastnumberoftreeandcropspeciesusedinagroforestry
and numerous pest species, a complete treatment of the subject matter is beyond the scope of this
chapter.Onlyaselectionofthemostwidelyusedagroforestrysystemsaregivenhereasexamples,
andtypicalcasesareexamined.Theobjectiveistoanalyzethefactorsthatinfluencepestincidencein
thelightofexistingecologicalhypotheses.Inthediscussions,moreemphasishasbeenoninforma-
tion generatedafter therecent reviews by Day and Murphy (1998),Rao et al.(2000),and Schroth
etal.(2000).Thisisintendedtofillthegapsinknowledgeandcomplementtheexistingreviews.
5.2 PARTITIONING THE COMPLEXITY OF PEST INTERACTIONS
In agroforestry systems, plants have close relations with abiotic and biotic components in the
community. According to Ong et al. (2004), the net effect of one plant component on another can
be expressedas:
I¼FþC þ Mþ PþL þ A,
where
Iis theoverallinteraction
Fiseffectson chemical, physical,and biological soil fertility
C iscompetition for light, water, andnutrients
Mis effect on microclimate
Piseffect on pests
L issoil conservation
Aisallelopathiceffects
Manyoftheseeffectsareinterdependentandcannotbeexperimentallyestimatedindependentlyof
oneanother.Thismeansthatwhenstudyingtheeffectofpestsinagroforestry,wecannotignorethe
effects inter alia of soil fertility, competition, or microclimate.
Pestsofanagroforestrysystemareessentiallythepestsofitscomponents(thecropsandwoody
perennials),andtheirdynamicsisgovernedbythecomplexityanddegreeofinteractionbetweenthe
crop, tree, and the composition of other plant communities such as weeds. Direct interactions
between treesand crops for growth resourcesmay exerciseastronginfluenceonpestsand natural
enemies of either or both components of the system (Table 5.1). In the following discussion, the
manner in which each component affects the other in terms of pest populations is briefly summ-
arized. A simplified model of potential interactions between the plant community, herbivores,
pathogens,andnaturalenemiesin a simultaneous agroforestry practice ispresented inFigure 5.1.
5.2.1 INTERACTIONS BETWEEN THE PLANT COMMUNITY, HERBIVORES, AND THEIR
NATURAL ENEMIES
The plant community (or producers), including the trees, crops, and weeds, constitute the first
trophic level. Each plant species may be attacked by a wide range of herbivores (i.e., primary
consumers), which constitute the second trophic level. Herbivorous species in turn are attacked
by natural enemies (i.e., secondary consumers), which constitute the third trophic level.
Natural enemies include predatory arthropods (e.g., insects, predaceous mites, spiders, scorpions,
centipedes, etc.) and vertebrates (e.g., insectivorous birds and mammals), parasitic insects (i.e.,
parasitoids), and pathogenic bacteria, viruses, fungi, protozoa, and nematodes, which play a
significantrole in thepopulation dynamics of pests of agroforestry (Sileshi et al.,2001).
Theinteractionsthatoccurbetweentheplants, herbivores,andtheirnaturalenemiesarecalled
tri-trophic interactions.The plant community may affect these interactions ina variety of ways, as
depictedinFigure5.1andTable5.1.Forinstance,treesthroughshadingortheirphysicalpresence
may directly influence the migration, host location, and feeding of insect pests of the crop in
Batishetal./EcologicalBasisofAgroforestry 43277_C005 PageProof page 76 25.6.2007 6:29pm CompositorName:BMani
76 EcologicalBasisofAgroforestry
TABLE 5.1
Summary of Tree–Crop Interactions and their Consequenceson Pests and Diseases
in Major Groups of Agroforestry Systems
Process PossibleEffects
Sequentialsystems Treecanopyshading=smotheringthe Reductionofannualandperennialweeds
understoryvegetation
Tree=shrubspeciesmaystimulategermination Weedseed-bankdepleted
ofparasiticweedStriga
Strigapopulationanditsseed-bankarereduced
Treesproducingallelopathicchemicals Reductionofweedpopulations
Treespeciesprofuselyproducingseedand Treespeciesbecomesanenvironmentalweeds
volunteerseedlings
Increasecostsofcontrol
Treeinfalloworboundaryplantingharboring Increasedpestsdamageinadjacentcropfields
pests
Increasesthepoolofavailablesoilnutrients, Increasedcropvigortowithstandsomepests
especiallyinorganicN
Increasedvigorinducingsusceptibilitytoother
pests
Treefallowsbreakingthecyclesofinsectand Reductionininsect,diseaseandnematode
pathogens damageonsubsequentcrops
Treesservingasalternativehoststoinsects, Increasedpestdamageonsubsequentcrops
nematodesandpathogens
Mulchesincreasingsoilhumidityandlowers Increasedsoil-bornediseasepopulations
soiltemperature
Treesservingasrefugeandfoodsourcefor Reductionofpestproblemsinadjacentcrop
naturalenemies fields
Simultaneoussystems Treesdominatingcropsbycompetitionfor Reducedvigorinducingsusceptibilitytopests
growthresources attack
Treesservingasrefugeandfoodsourcefor Reductionofpestproblemsinadjacentcrop
naturalenemies fields
Treeslinesactasmechanicalbarriersforthe Reductionofpestcolonization
spreadinsectpests,vectorsandpathogens
Treesimprovingmicroclimateinharsh Increasedcropvigor
environments
Buildupofpestsandpathogens
Treesservingasalternatehoststocroppests Increasedpestdamageoncrops
anddiseasevectors
Treepruningsusedasmulch Reductionofshadesensitiveweeds
Treeandcropsharingthesamepest Increaseinpestproblems
Treecanopyandleaflitterkeepingtheground Buildupofsomedisease
coveredformostpartoftheyear
additiontoactingasrefugefornaturalenemies.Treescanalsoinfluencepestincidencebyactingas
alternative hosts of acroppest orvector ofa pathogen. Trees, throughtheirindirect effectson the
nutrition of the crop, may also influence demographic parameters of crop pests such as natality,
longevity, and mortality. Thisinturn may trigger changes in themigration,host location, feeding,
and demographic patterns of natural enemies. Trees may also cause shading and reduce air
circulation, leading to high humidity and an increase in disease incidence. A detailed knowledge
of tri-trophic interactions associated with a given pest or pest complex is required if refuge for
naturalenemiesare tobe conserved orestablished.
Batishetal./EcologicalBasisofAgroforestry 43277_C005 PageProof page 77 25.6.2007 6:29pm CompositorName:BMani
Weeds,Diseases,InsectPests,andTri-TrophicInteractionsinTropicalAgroforestry 77
Natural
enemies
Crop harboring
Herbivores and vectors of tree
pathogens attacking diseases
woody perennials
Competition
Allelopathy
Shading/mulch
Addition of nutrients
Tree Alternative host of
Crop
crop pests
Refuge and food for
natural enemies
Herbivores and
pathogens
attacking crop
Competition
Competition Allelopathy
Shading Shading
Allelopathy Alternative host of Natural
Alternative crop pests enemies
hosts of tree Refuge and food for
pests natural enemies
Weeds
harboring Weeds Weed
vectors of harboring
tree diseases vectors of crop
diseases
Herbivores and
pathogens attacking
weeds
Herbivores act
as alternative
host of natural
enemies
Natural enemies
FIGURE5.1 Potentialinteractionsbetweentheplantcommunity,herbivores,pathogensandnaturalenemiesin
asimultaneousagroforestrypractice.
Weeds,inadditiontocompetingwiththetreeandcropcomponents,mayalsoactasalternative
hostsofpestsofthetreeorcropcomponents.Forinstance,inwesternKenya,Strigahermonthica,
aparasiticweedofcereals,isagoodhostforroot-knotnematodes,whichattackagroforestryspecies
such as Sesbania sesban and Tephrosia vogelii (Desaeger et al., 2004). Cultivated ground cover
plants and weeds (e.g., in orchards) can increase the heterogeneity of the habitat, alter the quality
and quantity ofbio-resources, and regulate ecological niches of various speciesinthe community.
Suchplantscanprovideavarietyofresourcesforpredatorsandparasitoids,includingshelter,food,
andinformationonthelocationoftheirherbivorousprey(BuggandWaddington,1994;Liangand
Batishetal./EcologicalBasisofAgroforestry 43277_C005 PageProof page 78 25.6.2007 6:29pm CompositorName:BMani
78 EcologicalBasisofAgroforestry
Huang, 1994). Liang and Huang (1994) reported that the weed Ageratum conyzoides, growing
in citrus orchards, plays an important role in stabilizing populations of the predatory mites
(Ambleyseius spp.), which are effective natural enemies of the citrus red mite (Panonychus citri).
Understoryvegetationcanalsosustainsignificantlyhighergeneralistpredatorssuchasladybeetles,
ground beetles, hover flies, mirid bugs, and lacewings in orchards than clean-weeded orchards
(Bugg and Waddington, 1994). Many aphids that colonize weeds can play an important role as
reservoirs of polyphagousnaturalenemiessuch as lady beetles,hover flies, and lacewings.
5.2.2 INTERACTIONS BETWEEN HERBIVORES AND PLANT PATHOGENS
The manner in which herbivores interact with plant pathogenic organisms include: (1) acting as
vectors, (2) wounding agents, (3) host modifiers, (4) rhizosphere modifiers, and (5) resistance
breakers(Agrios,1988).Desaegeretal.(2004)providespecificexamplesofsuchinteractions bet-
ween nematodes and soil-borne pathogens. Homopterous insects, beetles, and mites vector viral,
bacterial,andfungaldiseases,whichcausesubstantiallygreaterlossesthanthosecausedbythedirect
feedinginjurybytheinsects.Forinstance,thegreenpeachaphid(Myzuspersicae)isknowntobea
vector of more than 180 virus diseases. The cotton aphid (Aphis gossypii) transmits more than
80kindsofvirusdiseases.Theblackcitrusaphid(Toxopteracitricidus)isavectorofvirusdiseases
ofcoffee,citrustristezavirus,citrusinfectiousmottlingvirus,andlittleleafandlemon-ribbingvirus
of lemon (Michaud, 1998; EPPO, 2006). Some xylem fluid-feeding leafhoppers also transmit the
bacterial plant pathogen Xylella fastidiosa, which induces diseases of grapevines (e.g., Pierce’s
disease) and citrus (citrus variegated chlorosis), and also other diseases of coffee and stone fruits.
Citrus-variegatedchlorosistransmittedbytheglassy-wingedsharpshooter(Homalodiscacoagulata)
hasnowexpandedthroughoutmanycitrus-growingareasofSouthAmerica(Redaketal.,2004).
One of the classic examples of a disease vectored by beetles is the Dutch elm disease,
a vascular-wilt fungus, Ophiostoma (Ceratocystis) ulmi, carried from an infected tree to a healthy
onebybarkbeetlesofthegenusScolytus(Agrios,1988).Recently,theweevilPissodesnemorensis
has been reported as a vector of and wounding agent of the pitch canker fungus (Fusarium
circinatum)andDiplodiapineacausingdiebackonpines(Pinusspecies)(GebeyehuandWingfield,
2003).ThebeanbeetleOothecamutabilis,whichattacksSe.sesban,alsotransmitscowpeamosaic
virus,oneofthecommonestviraldiseasesofcowpeareducingyieldsbyupto95%(vanKammen
et al., 2001). Arthropods that transmit plant diseases may vector plant pathogens to and from the
tree, crop,and weed hosts inagroforestry (Figure5.1).
5.2.3 INTERACTIONS AMONG HERBIVORES
Interactionsalsooccuramongherbivoresintheformofcompetitionandmutualism.Competitionis
definedastheinteractionbetweenindividuals,broughtaboutbyasharedrequirementforaresource
in limited supply, and leading to a reduction in the survivorship, growth, and reproduction of the
competingindividuals(Speightetal.,1999).Generally,competitioncanoccuramongindividualsof
the same species (intraspecific) or members of different species (interspecific). Damage by one
herbivorespeciescouldinfluencepopulationsofasecondspeciesthroughchangesinplantquality,
even if the herbivores lived at different times of the year. West (1985) demonstrated that spring
defoliation by caterpillars of two Lepidoptera, Operophthera brumata (Geometridae) and Tortrix
viridana (Tortricidae), on oak leaves can reduce leaf nitrogen content, which adversely affects the
survival of the Lepidopteran leaf-miner Phyllonorycter (Gracillaridae) and aphids later in the
season.
Mutualismisatypeofsymbiosisinwhichtwoormoreorganismsfromdifferentspecieslivein
close proximity to one another and rely on one another for nutrients, protection, or other life
functions. For example, many ants are known to tend homopterous pests such as aphids, mealy
bugs,andscaleinsects,wheretheantsprotecttheseinsectsfrompredationandparasitism.Inturn,
Batishetal./EcologicalBasisofAgroforestry 43277_C005 PageProof page 79 25.6.2007 6:29pm CompositorName:BMani
Weeds,Diseases,InsectPests,andTri-TrophicInteractionsinTropicalAgroforestry 79
theantsgethoneydewfromtheirhosts.Ontheotherhand,antsarepredatorsandmaywellhavea
positiveeffectasbiocontrolagents.InshadecoffeeproductionsystemsVandermeerandcoworkers
(2002) demonstrated that ants (Azteca sp.) can have potential as pests through their positive effect
on scale insects, but also have potential as biological control agents through their effect on other
herbivores.
5.3 INTERACTIONS IN SELECTED AGROFORESTRY PRACTICES
Section 5.3.1 presents the characteristics of the various agroforestry practices as they affect the
occurrence and development of weeds, insect pests, and diseases. Agroforestry systems were
broadly grouped into sequential (rotational) and simultaneous systems (Rao et al., 1998). The
presentation was structured from the simplest to the more complex tree–crop associations to
facilitate comprehension of theinteractions.
5.3.1 SEQUENTIAL AGROFORESTRY PRACTICES
5.3.1.1 Rotational Woodlots and Improved Fallows
Intherotationalwoodlotsystem,foodcropsareintercroppedwithleguminoustreesduringthefirst
2–3 years. Then the trees are left to grow, harvested in about the fifth year, and food crops are
replanted (Otsyina et al., 1996). The food crops grown following the tree harvest are expected to
benefitfromimprovedsoilconditionsbythewoodlotspecies.Improvedfallows,ontheotherhand,
consistofdeliberatelyplantedspecies—usuallylegumeswiththeprimarypurposeoffixingnitrogen
as part of a crop–fallow rotation (Mafongoya et al., 1998; Sanchez, 1999). The legumes can be
planted as either single species or mixed stands. Compared with single-species fallows, mixed-
species fallows are believed to increase the biodiversity and sustainability of the fallow system,
provideinsurance against failure,produce multipleproducts,improve utilizationofavailable plant
growth resources,and reduce buildup of pests (Gathumbi,2000; Sileshi andMafongoya, 2002).
Raoetal.(1998)recognizedthreedistinctphasesbasedonthemajorsoilchangesthatoccurin
therotationoftreefallowsbycrops.Thesechangesmaydirectlyorindirectlyaffectthepopulations
ofweeds,pathogens,andinsectpestsaffectingthesubsequentcrop(Schrothetal.,2000;Sileshiand
Mafongoya,2002,2003).Oneofthesignificantimpactsofthesechangesinvegetationcoverison
the parasitic weeds (Striga spp.), which are widespread in most parts of sub-Saharan Africa and
causeannualcerealyieldlossesestimatedbetween$7and13billion(Annon,1997).Intwoseparate
studies conducted in eastern Zambia (Sileshi et al., 2006), rotational fallows of Se. sesban signi-
ficantly reduced incidence of Striga asiatica on subsequent maize compared with continuously
cropped monoculture maize, or that grown after a traditional bush fallow. This effect of the
Se. sesban fallow persisted through three consecutive cropping cycles. Similarly in Kenya,
Se.sesbanreducedthenumberofSt.hermonthicaseedsinthesoilby34%,whereasinmonoculture
maize plots the Striga populations increased over the same period by 11% (ICRAF, 1993). The
effect of Se. sesban on Striga was due to the combined effects of Se. sesban causing suicidal
germinationofSt.hermonthica(i.e.,a‘‘trapcrop’’effect)andimprovingsoilinorganicN,whichis
knownto be detrimental toStriga (Gacheru and Rao,1998).
Treefallowsalsoreducetheincidenceofweedsingeneralincludingtheperennialgrassessuch
asspeargrass(Imperatacylindrica)(Garrity,1997).InSriLanka,weedpopulationswerelowerby
42% and 54% in maize planted in improved fallow of Crotolaria juncea and Tithonia diversifolia
than in a natural fallow (Sangakkara et al., 2004). In Nigeria, 3 years of planted fallows of
DactyladeniabarteriinNigeriacaused36%decreaseintheweedseed-bankrelativetothecropped
field,whereasthesamedurationofbushfallowincreasedtheweedseed-bankby31%(Akobundu
and Ekeleme, 2002). Studies in Zambia (Sileshi and Mafongoya, 2003; Sileshi et al., 2006) have
demonstratedthatsomelegumefallowscanreducetheinfestationofmaizebyarableweeds.Inone
Batishetal./EcologicalBasisofAgroforestry 43277_C005 PageProof page 80 25.6.2007 6:29pm CompositorName:BMani
80 EcologicalBasisofAgroforestry
study(SileshiandMafongoya,2003),totalweedbiomassinmaizegrownafteranaturalfallowwas
six times higher than that grown after pure Se. sesban and pigeon pea fallows. The weed biomass
was correlated negatively with leaf litter indicating that the reduction is due to smothering of the
weedsthroughinitial suppression ofaboveground weed growth, and thethick mulch layer formed
by the leaf litter from the fallow trees subsequently depleting the weed seed-bank (Sileshi and
Mafongoya,2003).Manyfallowspeciesreleaseawiderangeofcompounds,commonlyreferredto
asallelochemicals,whichcaninhibitweedseedgerminationorreduceweedvigor.Legumecover-
crop residues in the course of decomposition release volatile organic compounds with potential
herbicidal properties (Gallagheret al.,1999).
Rotational fallows have also been shown to affect plant-parasitic nematodes that attack crops.
Somefallowspecies(e.g.,Sesbania,pigeonpea,Tephrosia,andAcacia)arehostsforplantparasitic
nematodes such as Meloidogyne and Pratylenchus spp. (Page and Bridge, 1993; Duponnois et al.,
1999;DesaegerandRao,2000).WiththeintroductionofSe.sesbanforsoilfertilityimprovementin
the tobacco-growing areas of southern Africa, the root-knot nematode problem became serious
(Karachi,1995;Shirimaetal.,2000).InTanzania,Meloidogyneinfectionwasconsistentlyhighest
whentobaccowasplantedafter2yearsSe.sesbanfallowscomparedwiththecropinrotationwith
natural fallow (Shirima et al., 2000). In a study conducted in western Kenya, Meloidogyne
infestationcaused52%–87%yieldreductioninbeans(Phaseolusvulgaris)plantedafterSe.sesban
(Desaeger and Rao, 2000). A Crotalaria agatiflora cover-crop increased root-lesion nematode
(Pratylenchus zeae) populations to levels that could limit maize growth, whereas it decreased
Meloidogyne incognita and Mel. javanica populations during the same time (Desaeger and Rao,
2000).Inanotherstudy(DesaegerandRao,2001),beancropthatfollowedmixed-speciesfallowsof
Se.sesbanþTe.vogeliihadincreasedroot-knotnematodedamagecomparedwithbeangrownafter
pure fallows of the respective species. On the contrary, bean crops that followed Se. sesban þ
Crotalaria grahamiana and Te. vogelii þ Cr. grahamiana did not experience yield losses. In a
separatestudyconductedinthesameareainwesternKenya(Kandjietal.,2003),beansgrewpoorly
whenplantedafterTe.vogeliiandCr.grahamianabecauseofhighincidenceofMeloidogynespp.
in the first cropping cycle. In the second and third cropping seasons, while the population of
Meloidogyne spp. decreased, spiral nematode (Scutellonema spp.) populations increased, which
caused heavy losses of beans and maize planted after the legume fallows (Kandji et al., 2003).
Studies by Kandji and coworkers (2001) found a positive correlation of Scutellonema populations
withexchangeablebasesinthesoil.Pratylenchuspopulationswerepositivelycorrelatedwithbulk
density,whereasMeloidogynepopulationswerecorrelatedwithclay,potassium,andorganiccarbon
contentofthesoil.Ontheotherhand,ParatrichordorusandXiphinemapopulationswerecorrelated
with calcium and soil bulk density(Kandji etal., 2001).
Rotational fallows also have significant effects on the incidence of insect pests of crop plants.
According to Rao et al. (2000), chaffer grubs, which destroy maize seedlings, increased in maize
plantedafterSe.sesbanfallowsinKenya.Snoutbeetles(Diaecoderussp.)thatbreedonSe.sesban,
pigeon pea, Cr. grahamiana, and Te. vogelii during the fallow phase attacked maize planted after
fallowswiththeseplantspeciesineasternZambia(SileshiandMafongoya,2003).Inanexperiment
involving pure fallows and mixtures of these legume species, the density of snout beetles was
significantly higher in maize planted after Se. sesban þ Cr. grahamiana compared with maize
planted after natural grass fallow. The population of beetles was significantly positively correlated
withtheamountofnitrateandtotalinorganicnitrogencontentofthesoilandcumulativelitterfall
underfallowspecies(SileshiandMafongoya,2003).BesideSe.sesbanbeinganalternativehostof
the beetle (Sileshi et al., 2000), its mixture with other legumes appeared to offer a favorable
environmentfor thesurvivalof thebeetles duringthe fallow phase.
In the same study in eastern Zambia, Sileshi and Mafongoya (2003) recorded lower termite
damage(%lodgedplants)onmaizeplantedafterTe.vogeliiþpigeonpea,Se.sesbanþpigeonpea,
and pure Se. sesban than maize grown after natural fallow. Monoculture maize grown after the
natural fallow had about 11 and 5 times more termite damage compared with maize grown after
Batishetal./EcologicalBasisofAgroforestry 43277_C005 PageProof page 81 25.6.2007 6:29pm CompositorName:BMani
Weeds,Diseases,InsectPests,andTri-TrophicInteractionsinTropicalAgroforestry 81
Te. vogelii þ pigeon pea and Se. sesban þ pigeon pea, respectively. The higher termite damage
recordedinthenaturalfallowwasapparentlyduetostresscausedbyweedcompetition.Inanother
study conductedatfour sites ineasternZambia,Sileshi andcoworkers(2005) found nodifference
between treatments in termite damage on maize plants after Te. vogelii, Tephrosia candida,
Se. sesban, and Crotalaria pawlonia, a traditional grass fallow, monoculture maize grown with
and without fertilizer. Though the differences were not statistically significant, maize planted after
Te.candidafallowshadconsistentlylowertermitedamagethanfullyfertilizedmonoculturemaize
at three out of the four sites. In western Kenya, incidence and damage due to groundnut hopper
(Hilda patruelis) increased on farms where Cr. grahamiana was planted as a rotational fallow
compared with new sites (Girma, 2002). The abundance of natural enemies and tri-trophic inter- AQ1
actionsinrotationalwoodlotsandimprovedfallowshasnotbeenstudied.Rotationalsystemsatthe
landscapelevelmaycreateamosaicoffallowedandcroppedplotsandhowsuchasituationaffects
pests needs tobe evaluated.
5.3.2 SIMULTANEOUS AGROFORESTRY PRACTICES
5.3.2.1 Trees on Cropland
Raoetal.(1998)recognizedthreedistinctcategoriesoftreesoncropland—scatteredtrees,boundary
planting,andintercroppingofannualcropsbetweenwidelyspacedrowsoftrees.Scatteredtreesin
cropland, often known as ‘‘parklands’’ are widespread traditional practices in the semiarid tropics.
ThebestknownonesarethoseinvolvingFaidherbia(Acacia)albida,Parkiabiglobosa,Vitellaria
paradoxa, Azadirachta indica in West Africa, and mango, Melia volkensii, Adansonia digitata,
Parinari curatellifolia, Acacia spp. in the semiarid parts of eastern and southern Africa. Trees in
these systems are rarely planted but are derived from natural regeneration and are protected by
farmers. In such a setup, a pest may be shared between the tree and the associated crop or the
adjacent vegetation and the resultant interactions may assume considerable significance. For
instance, fruit flies (Ceratitis spp.) and false codling moth (Cryptophlebia leucotreta) are one
such group of pests with a wide host range (De Meyer, 1998). The marula fly (Ceratitis cosyra)
and false codling moths attack fruits of Uapaca kirkiana and Pari. curatellifolia as well as
commercialfruitsincluding mango, guava, avocado, peach, andcitrus(Sileshi, unpublished data).
Trees in boundary planting and intercropping systems are deliberately planted and managed.
Boundary planting involves trees on farm and field boundaries, soil conservation structures, and
terracerisers.Intercroppingsystemsusewidelyspacedrowsoffast-growingtreessuchasCedrela
odorata,Se.sesban,andGrevillearobustainbananaandbeanfields.Themanagementoftreesused
aswindbreaksaroundorchardsandsurroundingtreesandbusheshasalsoasignificanteffectonthe
populationsofpestorganismsandnaturalenemies.Theeffectoftreesoncroplandonpestshasbeen
reviewedbyRaoetal.(2000)andSchrothetal.(2000).However,systematicstudiesinvestigating
theeffect oftrees on cropland ontri-trophic interactions are virtually lacking.
5.3.2.2 MixedIntercropping
Mixed intercropping involves relay intercropping and coppicing legume fallows. In the context of
using leguminous trees for soil fertility replenishment, relay intercropping has been found to be
moreappropriatethanrotationalfallowsinareascharacterizedbyhighpopulationdensityandland
scarcity,wherefarmerscannotforegocropsforthetree–fallowphase.Atypicalsituationisthatof
southern Malawi where trees or shrubs such as pigeon pea, Tephrosia spp., and Se. sesban are
planted betweenrowsor withinthe rows ofan already establishedmaizecrop(Phiriet al.,1999).
Coppicing tree fallows are another variant of mixed intercropping combining the elements of
rotational fallow (the fallow phase) and intercropping (the resprouting phase) (Sileshi and Mafon-
goya,2006).Treespeciesthatresproutwhencutatfallowterminationarecalledcoppicingspecies.
The legume species used in coppicing fallows include Acacia spp., Gliricidia sepium, Leucaena
Batishetal./EcologicalBasisofAgroforestry 43277_C005 PageProof page 82 25.6.2007 6:29pm CompositorName:BMani
82 EcologicalBasisofAgroforestry
spp., Calliandra calothyrsus, Senna siamea, and Flemingia macrophylla. Pure stands of these
species are normally planted at a spacing of 131 m and the fallows are left to grow for 2–3
years.Attheendofthefallows,thetreesarecut,andtheleavesandtwigsareincorporatedintothe
soil witha hand hoe. Every time the stumps resprout,the coppice biomass iscut and incorporated
into the soil. Acereal crop, often maize, isplanted on theridges between thetree stumps.
Like the short-duration fallow species, legumes grown in mixed intercropping have a signifi-
cant impact on witch weeds. The incidence of St. asiatica was monitored (Sileshi et al., 2006) in
1995–1997 cropping seasons in coppicing fallows established in 1991 and 1992 at Msekera
in eastern Zambia. The density of St. asiatica weeds was lower in maize grown in the coppic-
ing fallows of S. siamea, Flemingia congesta, and L. leucocephala than in monoculture
maize, whereas maize grown in those of Ca. calothyrsus and G. sepium did not differ from
monoculture maize.
Legume trees grown in mixed intercropping can also influence insect pest populations. In a
study in Malawi, Sileshietal. (2000) found higher densities of the bean beetle (Ootheca benning-
seni) in farms where Se. sesban was relay cropped with legumes such as cow pea (Vigna
unguiculata), bean, soybean (Glycine max), and bambara groundnut (Vigna subterranea). In
another study in Zambia, the beetle density and damage was higher in farms where Se. sesban
was planted next to cowpea and Hyacinth bean (Dolichos lablab). The beetle caused 100%
defoliation ofboth Se. sesban and theother legumes (Sileshi etal., 2000).
Sileshiandcoworkers(2005)monitoredtermitedamageonmaizefor2yearsinanexperiment
establishedin1992(describedearlier)andasecondexperimentestablishedin1997atMsekera.In
the experiment established in 1992, maize grown in the traditional fallow and S. siamea had
significantly higher percentage of lodged plants than fully fertilized monoculture maize during
the 2001–2002 cropping season. The damage to maize grown in Ca. calothyrsus, G. sepium, and
Fl. macrophylla did not differ from that in monoculture maize. On the contrary, during the
2002–2003 cropping season, fully fertilized monoculture maize had significantly more damaged
plants than maize grown in the different fallows except Fl. macrophylla. In this experiment, total
inorganicnitrogen,soilwateratplanting,andcoppicebiomassappliedduringtheseasonaccounted
for 59%ofthe variance inthepercentage oflodged maize plants. Intheexperiment established in
1997,thepercentageoflodgedplantswassignificantlyhigherinfullyfertilizedmonoculturemaize
growncontinuouslywithoutfertilizerthaninmaizegrowninAc.anguistissimafallowsinthe2001–
2002croppingseason,whereasinthe2002–2003croppingseason,nodifferencewasnotedamong
treatments. The percentage of lodged maize plants was significantly correlated with preseason
inorganic nitrogen (Sileshi et al., 2005). Hardly any study investigated the effect of mixed inter-
cropping onnaturalenemies.
5.3.2.3 Alley Cropping
Alley cropping (also called hedgerow intercropping) involves continuous cultivation of annual
crops within hedgerows formed by leguminous trees and shrubs. The legumes are periodically
pruned and their biomass is applied either as mulch or incorporated into the soil to improve soil
fertility (Kang,1993).
Trees in alley-cropping arrangements can have significant effects on the incidence of weeds,
diseases, and insect pests. Studies in Kenya (Jama et al., 1991; Jama and Getahun, 1992) showed
42%–98% reduction in weed biomass in maize and green gram (Phaseolus aureus) alley cropped
with F. (Acacia) albida and L. leucocephala compared with the respective monocrops. In Costa
Rica, Rippin et al. (1994) reported a 52% and 28% reduction in weed biomass in maize grown
betweenErythrinapoeppigianaandG.sepiumhedgerows,respectively.Oneofthemostimportant
aspectsofalleycroppingiscontrolofproblematicweedssuchasspeargrass(Imperatacylindrical)
(Garrity, 1997). On Alfisols in Nigeria, hedgerows of L. leucocephala and G. sepium reduced the
population of speargrass by 51%–67%, aboveground biomass by 78%–81%, and belowground