WAGENINGEN UNIVERSITY PAPERS 2001-3

The use of plant material to protect stored leguminous seeds against seed beetles: a review

by S.J. Boeke, J.J.A. van Loon, A. van Huis, D.K. Kossou and M. Dicke

Laboratory of Entomology, Wageningen University, P.O. Box 8031, 6700EH Wageningen,The Netherlands Dateo f publication: 30-10-2001 A ia

BACKHUYS PUBLISHERS 2001

^r j EDITORIAL MESSAGE WAgeningen University Papers is the new title of Wageningen Agricultural University Papers, which series was published from 1988 upto 1999 (the last issue was 1999-4: Monopetalanthus exit, by J.J. Wieringa). In the year 2000 no issues were produced. Predeces­ sors of the journal were the Agricultural University Wageningen Papers (1984-1988) and the Mededelingen van de Landbouwhoge­ school (te) Wageningen (1918-1983). The publication, a series of irregularly published monographs (some­ times reviews or theses) mainly in the English language (with occasional French language issues), is a medium for larger-sized papers of Wageningen University staff members and their collabo­ rators, working in the fields of plant- and animal sciences. Until 1996 the series was funded centrally by the University, but because of financial considerations this policy has been abandoned. Since then, funding has been rather haphazard. The series has always been used as an official exchange medium for the Central Library of the University, which policy has also been abandoned. The exchange list has been reduced, and in the future the WAUP will be offered for exchange with library partners only on the basis of counter- performance. The Editorial Board is looking for ways to continue the publication of WAUP. Sincerely,

WAUP Editorial Board. Prof.dr.ir. L.J.G. van der Maesen, Chairman Dr.ir. I. Bos, Member Dr. W. Backhuys, Publisher

ISBN 90-5782-100-1

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Bibliotheek Planteziekfenkund::.] Centrum Binnenhaven 3 - Pootbus 8122 6700 ER WAGENINGEN Theus eo fplan tproduct st oprotec t storedseed s

THEUS E OFPLAN T PRODUCTS TOPROTEC T STORED LEGUMINOUS SEEDS AGAINST SEEDBEETLES :A REVIE W

Tableo f contents

Abstract 3

1 Introduction 5

2Powder s or fresh application 9 2.1 Developmental stage affected 19 2.1.1 Effects on adults 19 2.1.2 Effects onovipositio n 19 2.1.3 Effects onegg san dlarva e 20 2.2 Comparison ofbeetl e species 21 2.3 Comparison ofplan t species 21 2.4Duratio n ofth eeffect s 22 2.5 Effects onth e storedproduc t 23 2.6 Conclusion 23

3 Ash 25 3.1 Developmental stage affected 26 3.1.1 Effects on adults 26 3.1.2 Effects on oviposition 27 3.1.3 Effects on eggs and larvae 27 3.2 Comparison ofbeetl e species 27 3.3 Comparison ofplan t species 28 3.4Duratio n ofth eeffect s 28 3.5 Effects onth estore dproduc t 28 3.6 Conclusion 29

4 Volatile oils 31 4.1Developmenta l stage affected 35 4.1.1 Effects on adults 35 4.1.2Effect s on oviposition 36 4.1.3Effect s on eggsan d larvae 37 4.2 Comparison ofbeetl e species 37 4.3 Comparison ofplan t species 37 4.4Duratio n ofth e effects 38 4.5 Effects onth e storedproduc t 38 4.6 Conclusion 39

5Non-volatil eoil s 41 5.1 Developmental stage affected 50 5.1.1 Effects on adults 50 5.1.2 Effects on oviposition 51 5.1.3 Effects on eggs and larvae 51 5.2 Comparison ofbeetl e species 52 Tableo fcontent s

5.3 Comparison ofplan t species 52 5.4 Duration ofth e effects 53 5.5 Effects onth e stored product 54 5.6 Conclusion 54

6 Extracts 55 6.1 Developmental stage affected 61 6.1.1 Effects on adults 61 6.1.2 Effects on oviposition 61 6.1.3 Effects on eggs and larvae 62 6.2 Comparison ofbeetl e species 62 6.3 Comparison ofplan t species 62 6.4 Duration ofth e effects 63 6.5 Effects onth e storedproduc t 63 6.6 Conclusion 63

7 General discussion 65 7.1 Modeo f action 65 7.2 Comparison ofbeetl e species 65 7.3 Comparison of application modes 66 7.4 Duration ofth e effects 66 7.5 Effects onth e stored product 67 7.6 Conclusion 67

8 References 91 Theus eo fplan tproduct st oprotec tstore dseed s

THEUS E OFPLAN T PRODUCTS TOPROTEC T STORED LEGUMINOUS SEEDS AGAINST SEED BEETLES:A REVIE W

S.J. Boeke1,J.J.A .va n Loon1,A .va n Huis1,D.K . Kossou2 andM .Dicke 1 1: Laborator yo fEntomology , Wageningen University, P.O Box 8031, 6700E H Wageningen, The Netherlands 2:Facult é des Sciences Agronomiques,Universit éNational e du Bénin, 01B P 526 Cotonou Bénin

ABSTRACT Inmos t tropical regionsbeetle s attack stored seeds and constitute amajo r causeo f serious post-harvest crop losses.Beetle s deposit eggs onth e seed surface, the larvae develop insideth e seed and leavei t asadults .T oprotec t the storedproduc t against these insects,no t only synthetic insecticides,natura l enemies andphysica lmethod s canb e applied,bu t also formulations ofinsecticida l orinsect-repellen t plants.Thes emethod s eachhav ethen - advantages and their disadvantages. Aconsiderabl e body ofliteratur e has accumulated onplan t products asinsecticides ,wit hvariabl eresults .A n overview ofth e available literature onplan t products used against storagebeetle s (with afocu s onBruchida e infesting cowpea) is presented. Therevie w is structured according toth etyp e of formulation used to applyth eplan t product,i.e .powder , ash,volatil e oil,non-volatil e oilan d extract. Forplan t powders,preparatio n and application are easy andtherefor e manyplant shav e been used aspowder s inpilo t tests.Th equantitie s applied vary and theresult s from thetest s diverge considerably. Plant powders can affect all stages ofth edevelopin gbeetle . Inmos t cases,th e powders dono thav e anydirec t effects onth e stored product. Ash can offer aneffectiv e wayt oprotec t stored seeds against storage beetles, ifi t isapplie d in large quantities. The ashhinder s adultmovemen t and thushamper s oviposition. Disadvantages aretha t ash cannegativel y affect the appearance and germination ofth e stored seeds.Fo r small quantities of seeds, ashcoul d be achea p and safe alternative for chemical pesticides. Volatile oils are often effective but theyquickl y evaporate,unles s they are applied in airtight storage structures. The oils aremainl y effective against adultbeetle s either asrepellent s ora s toxicants,whic h causes oviposition tob e affected aswell .Du et o set-ups from whichbeetle s cannot escape,th erepellen t effect isofte n underestimated in laboratory tests.Applicatio n of volatile oils is easybu t should berepeate d for longter mprotection . Thevolatil e oil doesno t needt ob e indirec t contact withth e stored product, soth eproduc t isusuall y unaffected byth e treatment. The yield ofvolatil e oil from plants isgenerall y low.Distillatio n equipment anda lot ofplan t material areneede d to obtainth eoil . Non-volatile oilshav emainl yphysica l effects on theinsects .Egg s aremos t susceptible. They areeithe r asphyxiated orthei r attachment toth e seed surface and subsequent penetration ofth e hatched larva arehampered . Non-volatile oils aredifficul t to apply. The seedwil lb eprotecte d duet o thisphysica l effect onlyi fth e oil forms afilm coverin g itswhol e surface. The side effects of oilso nth e stored product arenumerous ,rangin g from achang e of colourt o an alteration oftast e and inhibition of seed germination. Toobtai n large enough quantities ofoil s largequantitie s ofplant s areneede d and alo t ofwork . Manyo fth e edible oils areusuall y for sale. Extracts of fresh or dryplant s areusuall y more effective thanpowders .The y aremostl y effective against adultbeetles , either asrepellent s or astoxicants . Other developmental stages ofth ebeetl e canb e susceptible aswell . Theeffec t of extractso nth e stored product is usually negligible. Large quantities ofplant s areofte n needed to obtain sufficient amounts of extracts. Solvent availability can causeproblems . Abstract

Manydifferen t plant specieshav ebee nteste d for their effect on thebeetles .I nthi s review,mor e than 400 species arementioned . However, the literature available doesno t allow a general ranking of insecticidal orrepellen t efficacy. This isdu et o the lack of experimental detail and statistical analyses ofth eresult s inman y ofth e cited publications. In addition, theefficac y of aplan t product depends on theplan t species, the individual plant, theplan t part and thetim e and way ofharvesting . Inbioassays , the application mode,th equantity , thepreparatio n ofth eplan t material,th e state ofth e product to store (e.g.intactnes s ofth e seed skin and/orpod ) and storage conditions (e.g. humidity, rateo f infestation) aswel l asth ebeetl e speciesteste d are important for the outcome ofth etests .Th emer e fact that natural products areuse d implies that considerable variation ist ob e expected inbioassa y outcomes. Indeed, literature results areofte n contradictory, notwithstanding the fact thatman yplan t species do show useful effects against seedbeetles . Themos t effective plants ormethod s of application areno t yetknown ,bu t results arepromisin g andplan t products canb e an effective replacement for chemical insecticides inth ebattl e against seedbeetles . Theus eo fplan tproduct st oprotec tstore dseed s

1 INTRODUCTION Cowpea (Vignaunguiculata (L.) Walp.)i sa n important crop intropica l regions, particularly inWes t and Central Africa. The green parts of theplan t areuse d asa vegetable or asfodde r for cattle.Th e seeds contain ahig h amount ofprotei n andB - vitamins (238) andhel p topreven t starvation among lowresourc e fanners and thepoo r urban population (80; 156;243) .Th eplant , in symbiosis with root nodule bacteria ofth e Rhizobiumgenus ,ca nfix nitroge n (225)an ddoe s notnee d extranitroge n or fertilisers toprovid e goodharvest s (15).Fo r many varieties,th enee d for water islimite d soth e plant can grow onpoo r and dry soils of, for example,th e Sahelian region. The spreading growth form ofth e plant covers the soil making it agoo d competitor against weeds in single orinter-croppin g systems(92) . Many diseases,viruse s and insect pests (41; 126) attack theplan t inth efield, bu t once the cowpea hasbee nharveste d theproblem s areno t over. After harvest, thebean sca n be either sold immediately for arelativel y lowpric e orthe y can be stored before useo r trading.A tth een do fth e storage season,jus tbefor e thene wharvest ,th emarke t price willb e much higher than immediately after harvest (305).However , by thattim e the storedproduc t can be damaged not onlyb y fungi androdent s but alsob y insects,i f nothing isdon et opreven tthis . Inth e field, several beetle species laythei r eggs onth e surface ofth e maturing pod or directly onth etest a ofth eripenin g seed andca nthu sb ebrough t toth e storage room with theharveste d beans (161; 343).Th e larvae ofthes e beetles develop insideth e bean, destroying itscontents .Afte r a few weeks,ne w adults emerge,leavin gth ebea na perforated seed with alo wprobabilit y of germination (38;307) .Thes ene w adults oviposit on the available beans again. Most ofth ebeetl e species with such alif e cycle have only few generations insideth e storage structures before they develop intoa diapausing (101)o rnon-reproducin g flying form, which leavesth e storageroo m(104) . However, onebeetl e species,th e cowpea weevil, Callosobruchus maculatusFabriciu s (Coleoptera:Bruchidae ) iscapabl e ofdevelopin g inside cowpea storehouses for many generations before going into diapause.Thi s beetle willno t develop intoth e flying form toleav e the storage structure until allbean sar ehollo w orpulverised . Theegg sar etake n from the cowpea field (105;241 ) into the storage roomwit hth e harvested beans. Females ofth ereproductivel y active,flightles s form canla yu pt o 120egg sdurin gth e 3-15day s of their adult life. The eggshatc h after 3-8 days andth e larvae develop in9 - 19day s insideth ebea n to thepupa l stage of 5-11days .Th ewhol e life cycle,fro m adult to adult, takes 21t o4 0 days,dependin g onth etemperature , humidity,hos t plant, etc. (88; 112; 197;221 ;313) .Thi sbeetl e isknow n to infest 100%o fa cowpea harvest within 3t o 5month s of storage(300 ) andi t isresponsibl e for over 90%o f all insect damage to cowpea seeds(44) . Due to insect infestation, the germination ofth e seeds decreases, (31; 147)an d incidence of fungal infection increases (46).Insec t infestation causes the uric acid content (310)an dth e contents ofanti-nutritiona l factors, such asphyti c acid, trypsin inhibitor activity and saponin ofth ebean st o increase (193),whil e theweigh t ofth e storedproduc t decreases (192).Th econtent s ofvitamin s ofth e B complex (thiamine, riboflavin andniacin ) (190)an do f starch,energ y andnon-reducin g sugarsdecreas e for stored seedswit h increasing infestation, whilecontent s ofcrud efibre, cellulose , hemicellulose and lignin increase (189).Th eprotei n quality changes with infestation, withprobabl y a decrease inmethionin econten t anda n increase ofnon-protei n nitrogen. Insect-infested legumes become unhygienic duet opresenc e ofhig h amounts ofuri c acidsan dar eno t suitable for human consumption (84; 191). 1 Introduction

Control measures against storage beetles There areman ymethod s known topreven t orreduc e damage done by the cowpea weevil.Th e most common control measures taken against storage beetles are: - Synthetic pesticides. Some chemicals haveprove n tob e very effective against bruchid damage ifthe y areuse d atth erigh ttime ,i nth eright quantities , with the rightapplicatio n method,etc .Fo r lowresourc e families invillages ,however , the availability and costs of such chemicals can bring about great problems whereasa lacko fknowledg e about the application mayreduc e theefficac y ofth epesticid e and can bea hazar d to consumers ofth ebeans . Moreover, resistance ofth ebeetle s to somepesticide s has already beenreporte d (28; 90;292) .Othe r disadvantages areth e fact that someo fthes eproduct shav ea severenegativ e effect on seedviabilit y (254) andtha t they kill all insects, including beneficial ones such asth enatura l enemies of thebeetles . - Hermetic storage. Storagebeetle s canpenetrat e plastics upt o 0.18m mthic k (297) but thick plastic bags can beuse d to storebean s (301) especially ifthe y areuse d with acotto n innerlinin g(43) .Oi l drumswit hwell-fitte d lids(286 )ar eals o practicable to store seeds ifthe y arefilled completely , with as littleai r left as possible.Th edevelopin g beetles will thenus eu pth e available oxygen (304) within a short time (ca.2 weeks) andthu s suffocate before their development can cause serious damage (43; 289).However , any tinyhol e inth eba g ordru m through which a flow ofai rca npas swil lnullif y the effect, sobag s shouldb eprotecte d against rodents,vessel s should beprotecte dfrom rus t andth e structures should be treated with care to avoid damage. - Natural enemies.Developin g bruchid beetles canb eparasitise d by egg parasites such as Uscana lariophaga orb y the larval orpupa lparasite s such asDinarmus basalisan dEupelmus vuilleti. Underoptimise d laboratory conditions, parasitization can control thebruchi d infestation up to 70%(107 ) or even upt o 82%(53) .I nth e field,th eparasitoid s also suppress thebuil d up ofbeetl e populations (43; 269; 273; 335; 337),bu t the control isneve r 100%.Fo r an overview, seeGahuka r(95) . - Inert materials. Sand or ashca n bemixe d with stored beanst omak e an effective barrieragains t thebeetle swhic hprevent s theemerge d adultsfrom finding eac h other for mating orfrom reachin g anex t bean to oviposit on.Fin e ash or sand can effectively suffocate the adults,larva e andpossibl y eggs (49).Th e large quantities ofiner t material needed makethi smetho d ofprotectio n less practical, especially for considerable quantities ofstore d beans. Physical methods. In a(solar )heater ,o ri nplasti c bags exposed toth e sun(48) , where atemperatur e ofa t least 57 °Cca n bereache d for more than onehour , all stages ofth ebruchi d arekille d whereas thecookin g properties and germination of thebean s are not negatively influenced (152).A t 51.5 °Cfo r 15minute s or at47. 5 °C for four hours all adult female beetles are killed (122).Youn g developmental stageso fth ebeetle s aremos t susceptiblet o such ahea t treatment (150). This treatment inhibits thetransformatio n from pupa to adults and induces adult mortality (276). Coldtreatmen t or freezing thebean s kills all developmental stages. Seedsca n alsob e preserved by hangingthe m inth e smoke ofa cookin g fire (153).I fcar e is taken topreven t re-infestation, these couldb eusefu l optionst opreven t damage. Sievingth ebean swil lremov e manyo fth eadul tbeetle s andthus prevent s severe attack (309). Gamma radiation is lethal tobruchids .Egg s and young larvae are especially sensitive tothi s treatment (102)an d females appear tob e more sensitive to it than males(82) . Theus eo fplan tproduct st oprotec t storedseed s

Treatment withbioga s (196;326 ) lowoxyge n and high carbon dioxide kills adult storage beetles and, lesseffectively , eggs,larva e andpupa e (115; 220)wit h pupae being least susceptible (179).A n anoxic atmosphere with 100%carbo n dioxide is more toxic to eggs,larva e andpupa e than atmospheres containing low levelso f oxygen (177). - Plant material. Manyplant s are known torepe l insects and toproduc e compounds toxict onon-specialise d insects.I fproduct s of suchplants , fresh ordrie d material, extracts or oils areapplie d to stored beans,the y havebee n shown to effectively protect stored cowpeas against bruchid infestation.

Plant products could offer asolutio n for theproblem s ofavailability , healthrisks ,costs , andresistanc e inth e caseo f synthetic pesticides,an d for the lack ofequipmen t for hermetic storage,gamm a irradiation and controlled atmospheres. They couldb e compatible with natural enemiest o reduce thepes tpopulatio n inth e seeds orthe y could replace space-consuming inert materials.I nth e ideal situation,plan t products couldb e readily available for everybody without costs,easil y andsafel y applicable andtoxi co r repellent only toth etarge t organism. Especially the lowresourc e farmers inth etropics , who can not afford chemical insecticides,wil lprofi t from cheapway s toprotec t their stored seeds. The effect and theefficac y ofth eplan t materials depend on their mode of application. Thematerial s can be added to thebean s asfres h wholeplants , or othermethod s canb e used to makethe mmor e effective. Upunti l now,muc h research hasbee n donet otes tman yplan t species for their efficacy assee dprotectants . However, nogenera l testprotocol s havebee n used and the applied quantities vary greatly. Many authors have tested plant materials andhav ereporte d their experiences,bu tn o overview exists ofth erankin g for insecticidal properties ofplants .

Apart from cowpea, many other seed crops such asmaize ,cereals ,peanuts , grams and beans, are also damaged during storage,b ybeetl e specieswit h life histories similar to that ofth e cowpea beetle. The methodsuse d against these insects are generally comparable tothos e employed against Callosobruchus maculatus.I n this paper knowledge ofcontro l measures usingplan t products against the storage beetle species mentioned intabl e 1,i sreviewe d ingeneral ,wit h afocu s onbeetle s ofth e family Bruchidae. The abbreviations given inthi s table areuse dthroughou t thepaper . Per application mode alis t ofplant s isgiven ,ordere d perplan t family, anddiscusse d with respect topossibl e correspondence or discrepancies. Inthi s discussion,th e differences in set-ups ofth etest s asthe y areuse d in different publications could mostly not be taken into account. 1 Introduction

Table1: Names, families, descriptorsand common namesof the beetle species mentionedin this article (After Couilloud 54). Beetle species Author(s) Beetle family Common name Abbreviation 1 Acanthoscelides obtectus Say Bruchidae Bean bruchid Ao 2 Attagenus megatoma Piceus (Olivier) Dermestidae Black carpet beetle Am 3 Bruchidius atrolineatus Piceus Bruchidae Pulse beetle Ba 4 Callosobruchus analis Fabricius Bruchidae Ca 5 Callosobruchus chinensis Linnaeus Bruchidae Adzuki bean beetle Cc 6 Callosobruchus maculatus Fabricius Bruchidae Cowpea beetle Cm 7 Callosobruchus phaseoli Gyllenhal Bruchidae Bean weevil Cph 8 Callosobruchus rhodesianus Piceus Bruchidae Cr 9 Callosobruchus subinnotatus Piceus Bruchidae Csu 10 Caryedon serratus Olivier Bruchidae Groundnut seed beetle Cse 11 Cylas puncticollis Boheman Apionidae Sweet potato weevil Cpu 12 Dermestes maculatus De Geer Dermestidae Common hide beetle Dm 13 Lasioderma serricorne Fabricius Anobiidae Cigarette beetle Ls 14 Oryzaephilus surinamensis Linnaeus Silvanidae Saw toothed grain beetle Os 15 Prostephanus truncatus Horn Bostrichidae Greater/larger grain borer Pt 16 Rhyzopertha dominica Fabricius Bostrichidae Australian wheat weevil Rd 17 Sitophilus granarius Linnaeus Curculionidae Grain weevil Sg 18 Sitophilus oryzae Linnaeus Curculionidae Rice weevil So 19 Sitophilus zeamais Motschulsky Curculionidae Maize weevil Sz 20 Tribolium castaneum Herbst Tenebrionidae Red flour beetle Tea 21 Tribolium confusum Jaquelin du Val Tenebrionidae Confused flour beetle Tco 22 Trogoderma granarium Everts Dermestidae Khapra beetle Tg 23 Zabrotes subfasciatus Boheman Bruchidae Mexican bean beetle 2s Theus eo fplan tproduct st oprotec tstore dseed s

2POWDER S OR FRESH APPLICATION The simplest wayt oappl yplant st o astoc k of seedsi sharvestin g theplan t andaddin g it toth e seeds.A sth e stored seeds should be dry to prevent moulds and germination, the plants areofte n pre-dried. Toattai n afiner an dmor eeve n distribution, these dryplant s can than be ground topowde r before application. Many plants have been tested inth e laboratory aspowder s toestimat e theirpossibl e effects. Themode s ofactio n ofthes e powders vary,bu t with lowt o moderate dosages,th e effect isalway srepellen t ortoxic , never mechanical. Allplant s discussed inthi s chapter and in Table 2wer e dried, ground anduse d aspowders ,unles s stated otherwise.

Table 2:Plants usedagainst storage insects, the quantityand the used plant part. Plant (sub-) Plant species Plant part $ Quantity Beetle Affected stage # Refe­ A family species A O E H L M P rence

Acanthaceae Adhatoda spp. L 8% Cm ** 111

Acanthaceae Adhatoda vascia L 5-20 Cc *! ** 1 L 5-20 Cc ** 228 Amaranthaceae Achyranthes aspera - 5-20 Cc *! *! 50 Amaryllidaceae Crinum defixum ~ 5-20 Cc *! *! 50

Anacardiaceae Anacardium occidentale Nut shell 4 perforatec Cm *! M *! 81 liquid nuts/kg Gum exudate 5-60% Cm *! *! *! 174 Anacardiaceae Sclerocarya birrea Bs 100-300 Cm *! 93

Annonaceae Annona reticulata L 0.1-0.4g / Cm ** *| *! 248 50 seeds Annonaceae Annona senegalensis L po/wh 1/10 pods Ba.Cm *• •* ** ** 15 Bs 100-300 Cm ** 93 Annonaceae Annona spp. S 0.5-2% Cm *! 111 Annonaceae Annona squamosa S 50 Ca ;; H M 135 S 5-100 Ca *i *! 139 S 10-50 Cc *! *! *! 296 Annonaceae Dennettia tripetala F 4-120 Cm.Sz n n 215 Annonaceae Monodora myristica L 100-300 Cc, Cm, Cr *! *! 251 F 4-60 Cm, Sz *i n 215 Annonaceae Xylopia aethiopica F 2 g/500 Cm ** ** 209 seeds F 4-60 Cm.Sz *i *! 215

Apiaceae Anethum graveolens S 5-20 Cm, Ls, So, *i 319 Tco 2Powder so rfresh applicatio n

Plant Family Plant species Plant part Quantity Beetle A O E H L M P Ref.

Apocynaceae Nerium oleander L 60 Cm *! *! ** 131 L,T 40 Cm *! 341 Apocynaceae Thevetia peruviana F 5-20 Cc *i 228 F 1-2% Cm *! 111 Araceae Acorus calamus - 5-20 Cc H *! 50 R 5-20 Cc *! 228 R 10-50 Cc *! *! *! 296 T 0.5-5% Cc, Sg, So *! 121 R 1-2% Cm *! 111

Asteraceae Ageratum conyzoides T 2 g/500 Cm ** ** 209 seeds Asteraceae Artemisia tridentata L,T 1-30 So, Zs *! *! *! 348 Asteraceae Aspilia africana T 2 g/500 Cm ** ** 209 seeds Asteraceae Balsamorhiza sagittata L,T 1-30 So,Zs ** *! *! 348 Asteraceae Chromolaena odorata L 0.5-4 g/50 Cm ** ** 2 seeds L 12.5-75 Cm *! 12 L 2 g/500 Cm ** ** 209 seeds G 25 Cse ** *! *! 67 Asteraceae Emilia sonchifolia L 2 g/500 Cm ** ** 209 seeds Asteraceae Eupatorium spp. Wh 100 Ao ** 30 Asteraceae Sphaeranthus indicus L po/wh 50 Cc *! 325

Cactaceae Opuntia burrageana L 100-300 Cc, Cm, Cr *! *! 251

Capparaceae Boscia senegalensis L po/wh Layers Ba, Cm *! *! *! 13 L po/wh 3/10pod s Ba, Cm *i ** *! *! 15 Twh Layers Cm ** 59 - 25 Cm *i ** ** 60 F, L fr/d 0.5-32% Cm ;i !l M 288 F fr 1.2-4.8g/ 1 Cm M 289 F,L fr 20-40 Cm, Cse, Pt, *! *! *i 180 Sz, Tea

Chenopodiaceae Chenopodium G 25 Cse *! H *! 67 ambrosioides

10 Theus eo fplan tproduct st oprotec tstore dseed s

Plant Family Plant species Plant part Quantity Beetle A O E H L M P Ref.

Combretaceae Combretum apiculatum B,L 60 Cm *! *! ** 131 Combretaceae Combretum imberbe Wood 60 Cm *! *! ** 131 Combretaceae Terminalia sericea B,L 60 Cm *! *i ** 131

Convolvulaceae Ipomoea carnea L 5-20 Cc *! 228 L 8% Cm *i 111 Convolvulaceae Ipomoea mauritiana L 10-50 Cm ** ** 69

Cucurbitaceae Momordica spp. L 2g/500 Cm ** ** 209 seeds

Dilleniaceae Dillenia retusa L 0.1-0.4 g / Cm *! *! *! 248 50 seeds

Ericaceae Ledum palustre T 0.5-5% Cc, Sg, So *! 121

Euphorbiaceae Bridelia ferruginea B 2g/500 Cm ** ** 209 seeds Euphorbiaceae Croton gratissimus L 60 Cm *! *! ** 131 Euphorbiaceae Euphorbia tirucalli T 40 Cm *! 341 Euphorbiaceae Jatropha indica L 20 Cm, Sz ** 22 Euphorbiaceae Ricinus communis L 3.3-16.7 Cm u *! *! 214 Euphorbiaceae Spirostachys africana B 60 Cm *! *! ** 131

Geraniaceae Geranium viscosissimum L,T 1-30 So.Zs *! *! *i 348

Juglandaceae Juglans spp. Nut shell lg/60 Ao ** 51 insects

Lamiaceae Hyptis spicigera Wh 0.3-30 Ao ** *! 162 whfr 3-5 cm Ba, Cm ** 58 layers L Layers Cm ** ** ** ** 59 -- 25 Cm *! ** ** 60 L, T 33.3 Cm, Rd, So, *! 8 Tea, Tg Lamiaceae Hyptis suaveolens Shoot 100-300 Cm *! 93 T 25 Cm, Rd, So, ** 8 Tea, Tg Lamiaceae Lavandula angustifolia -- 1-7.4 Ao.Sg *! 138 Lamiaceae Mentha piperita Wh 1 g/6 beans Ao *! ** ** ** 261 L 50 Ca M n M 135 Lamiaceae Mentha spicata L 10% Ca *! *! 100

11 2Powder so rfresh applicatio n

Plant Family Plant species Plant part Quantity Beetle A O E H L M P Ref. Lamiaceae Monarda fistulosa L,T 1-30 So.Zs *! *! »! 348

Lamiaceae Ocimum basilicum L lg Ao, Zs !! 344 L 10-50 Cc ** ** ** 296 Wh Layers Cm ** 59 - 25 Cm *! ** ** 60 G 25 Cse ** ** ** 67 L 5-50 Zs *! Ü *! *! 346 Lamiaceae Ocimum gratissimum Wh Layers Cm ** 59 -- 25 Cm *! ** ** 60 T 2g/500 Cm *! *! 209 seeds Lamiaceae Ocimum sanctum L 0.1-0.4 g / Cm *i *! *! 248 50 seeds Lamiaceae Origanum vulgare - 1-7.4 Ao.Sg *! 138 Lamiaceae Rosmarinus officinalis Wh 1 g/6 beans Ao *! *! *! *! 261 - 1-7.4 Ao.Sg *! 138 Lamiaceae Satureja hortensis wh 1 g/6 beans Ao *i *! *! *! 261 Lamiaceae Tetradenia riparia L 1-100 Ao, Zs *! *! *! 345 Lamiaceae Thymus serpyllum wh 1 g/6 beans Ao *! *! *! *! 261 Lamiaceae Thymus vulgaris wh 1 g/6 beans Ao *! *! *! *! 261 -- 1-7.4 Ao.Sg *i 138

Lauraceae Cinnamomum camphora L 100-300 Cc, Cm, Cr *! *! 251 L 10-50 Cm ** ** 69 Lauraceae Laurus nobilis wh 1 g/6 beans Ao ** ** *! ** 261 -- 1-7.4 Ao.Sg *i 138

Lecythidaceae Napoleona imperialis L 5-37.5 Cm, Csu, Sz ** ** ** 176

Leguminosae- Chamaecrista nigricans -- 0.003 Ao *! 66 Caesalpinioideae wh 0.3-30 Ao ** *! 162 L 2g/20seeds Cm ** *! ** ** 59 - 25 Cm *i *! ** 60 - 10ml/k g Cm ** 61 L, P 50 ml/1 Cm, Tco *! 354 Leguminosae- Erylhrophleum B 2g/500 Cm *! *! 209 Caesalpinioideae suaveolens seeds Leguminosae- Peltophorum africanum B 60 Cm *! ** ** 131 Caesalpinioideae

12 Theus eo fplan tproduct st oprotec tstore dseed s

Plant Family Plant species Plant part Quantity Beetle A O E H L M P Ref. Leguminosae- Senna occidentalis L, S 100 Cm ** ** 165 Caesalpinioideae Leguminosae- Tamarindus indica L 10-50 Cm *! *! 69 Caesalpinioideae

Leguminosae- Acacia concinna Pod 5-10 Cc *! ** 1 Mimosoideae Leguminosae- Acacia sinuata L,S 10% Ca ** ** 100 Mimosoideae S 10-50 Cc »* ** ** 296 Leguminosae- Entada africana L 100-300 Cm ** 93 Mimosoideae Leguminosae- Parkia biglobosa Bs 100-300 Cm ** 93 Mimosoideae Leguminosae- Prosopis africana Bs 100-300 Cm *i 93 Mimosoideae

Leguminosae- Derris inudata L 100-300 Cc, Cm, Cr *! *! 251 Papilionoideae Leguminosae- Derris spp. 3% Cc, Sz *! 200 Papilionoideae Leguminosae- Myroxylon balsamum L 10-50 Cm ** ** 69 Papilionoideae Leguminosae- Phaseolus vulgaris Beans' 0.1-5% Cm *! 130 Papilionoideae phytohemag glutinin Leguminosae- Pongamia pinnata S 10-50 Cc ** ** ** 296 Papilionoideae L po/wh 50 Cc *t 325 Leguminosae- Tephrosia vogelii G 25 Cse *i M H 67 Papilionoideae L 0.8-80 So.Zs *i 207 Leguminosae- Trigonella foenum- L, S 100-400 Ao, Tea *! *! *! *! 234 Papilionoideae graecum

Liliaceae Allium sativum Cloves fr 1 g/6 beans Ao ** ** ** ** 261 Bu 20 Cm ** ** 132 Liliaceae Aloe marlothii L 60 Cm *! *! ** 131

Malvaceae Sida acuta T 2g/500 Cm *• ** 209 seeds

Meliaceae Aphanamixis polystachya B,L,S 25 Cc *! *! ** *! *! 330

13 2 Powders or fresh application

Plant Family Plant species Plant part Quantity Beetle A O E H L M P Ref. Meliaceae Azadirachta indica L 0.25-11/ 1 Ao, Zs *! 49 L po/wh 1/10pod s Ba, Cm ** ** *» ** 15 K,L 10% Ca ** *» 100 S 50 Ca il M II 135 K,L 5-20 Cc *! *! 1 L,S 1-3% Cc il ii ii 4 K 3% Cc ** 40 - 5-20 Cc *! *! il 50 L 2-8 Cc *! 226 B 2-8 Cc *! 227 L 20-80 Cc *! *! 267 S 10-50 Cc *! *! *! 296 - 2-8 Cc *! 302 w 2-8 Cc *! 308 L po/wh 50 Cc *! 325 K, L po/wh 0.5-15% Cc, Cm. Rd, *! *! *! *! 275 Sg, So, Sz, Tea, Tco, Tg K,L 0.5-5% Cc, Sg, So *! 121 K,L 50 Cm *| 10 F 12.5-75 Cm *! 12 L 2g/20seeds Cm ** ** ** ** 59 K,L 25 Cm *! *• ** 60 S 10ml/k g Cm ** ** 61 F 5 Cm *! *! *! 81 L,K, S 10-20,0.5 - Cm *! *! 111 12% K 50-150 Cm ** *i ü |t 123 K 5-20 Cm M 134 S 6.4-38.4 Cm *! *! 161 S 0.5-5% Cm !! *! *! *! 169 L 0.5-3% Cm il *! u M 170 S 25 Cm *! *! 211 S 6.25-150 Cm *! *! H 212 L,S 10-30 Cm *! 285 L,S 10-30 Cm *! ** *! 287 K 0.5-2% Cm *! *! 311 L 20 Cm.Sz H 22

14 Theus eo fplan tproduct st oprotec t storedseed s

Plant Family Plant species Plant part Quantity Beetle A O E H L M P Ref. Meliaceae Khaya senegalensis B 2g/20seeds Cm ** »* ** ** 59 Bs 100-300 Cm ** 93 Meliaceae Melia azedarach -- 50 Cm *i 10 F 1-8% Cm *! *! 85 F,B 60 Cm *i »! ** 131

Moraceae Ficus exasperata L 2 g/500 Cm »» ** 209 seeds

Myrtaceae Eucalyptus citriodora L po/wh 20-80 Ce M 325 Myrtaceae Eucalyptus globulus L po/wh 1 g/6 beans Ao ** *! *! *! 261 Myrtaceae Eucalyptus spp. L 10-50 Ce ** ** ** 296 L po/wh 50 Ce *! 325 F,L 60 Cm *! *! ** 131 L 20 Cm.Sz ** 22 Myrtaceae Eugenia aromatica Fl 20 Cm n M 132 Myrtaceae Eugenia uniflora L 0.5-4 g/50 Cm *! *! 2 seeds

Piperaceae Piper guineense F po/wh 25-75 Cm *! *! *! 125 S 12.5-37.5 Cm *! il M 178 F 53-420 Cm *! *! 216 F 5-37.5 Cm, Csu, Sz *! *! *! 176 F 4-120 Cm.Sz M ü 215 Piperaceae Piper nigrum -- 0.012 Ao *i 30 S 2.8-11.1 Ao *! 163 S 0.03-0.12% Ao, Ce *! *! M 199 S 50 Ca H n n 135 -- 0.05% Ca, Cc ** 149 -- 0.25-0.5% Cc M 111 - 0.12% Cc *! 198 s 10-50 Cc *! *! *! 296 F 20 Cm *! *i 132 L 0.10 g/50 Cm *! 247 seeds L 0.1-0.4g / Cm *! *! *! 248 50 seeds F 20 Cm *! 341 F 0.063-0.5% Cm, So *! *! 315 — 10 g Cm, Sz *! 22

15 2 Powders or fresh application

Plant Family Plant species Plant part Quantity Beetle A O E H L M P Ref.

Poaceae Cymbopogon citratus L 100-300 Cc, Cm, Cr *! *! 251 L 0.5-4 g/50 Cm ** ** 2 seeds L 10-50 Cm ** ** 69 L 2g/500 Cm ** *! 209 seeds L,T 40 Cm ** 341 Poaceae Cymbopogon nardus wh 1 g/6 beans Ao ** *! *! *! 261 Poaceae Cymbopogon B 2g/20 Cm ** *! ** ** 59 schoenanthus seeds L 25 Cm *! ** •* 60

Polygalaceae Polygala butyracea Seed coat 250 ml/1 Cm, Tco *! 354 Polygalaceae Securidaca L, Br 50-100 Cm, Sz, Tea *! *! *! 180 longepedunculata

Polygonaceae Polygonum hydropiper L 20-80 Cc *! *! 267

Rubiaceae Diodia sarmentosa Fl 5-37.5 Cm, Csu, Sz ** ** ** 176 Rubiaceae Mitracarpus scaber Shoots 100-300 Cm ** 93 Fl 5-37.5 Cm, Csu, Sz ** ** ** 176

Rutaceae Citrus aurantifolia L 100-300 Cc, Cm, Cr ** ** 251 P 50-100 Cm ** ** *! 219 Rutaceae Citrus crematifolia P 0.20 g/50 Cm *! 247 seeds P 0.1-0.4g / Cm ** *! *! 248 50 seeds Rutaceae Citrus grandis P 40 Cm *! 341 Rutaceae Citrus limon L 100-300 Cc, Cm, Cr ** ** 251 P 50-100 Cm ** ** ** 219 L,P 20 Cm *! 341 Rutaceae Citrus paradisi P 1-100 Cm it *! H 9 P 50-100 Cm *i *! *! 219 P 25-180 Cm, Dm *! 73 Rutaceae Citrus reticulata P 50 Ca !! u M 135 P 40 Cm ** 341

16 Theus eo fplan tproduct st oprotec tstore dseed s

Plant Family Plant species Plant part Quantity Beetle A O E H L M P Ref. Rutaceae Citrus sinensis L 100-300 Cc, Cm, Cr *! *! 251 P 1-100 Cm *i *! *i 9 P 12.5-75 Cm *! 12 P 2-4 Cm ** 37 L 10-50 Cm ** ** 69 P 50-100 Cm ** ** ** 219 P 25-180 Cm, Dm *! *! 73 P 20 Cm, Sz ** 22 Rutaceae Citrus spp. Fwh 1/5 kg Ao ** 30 Rutaceae Zanthoxylum Br, S 25 Cm *! *! 211 zanthoxyloides Br 6.25-150 Cm *! *! M 212 Br 25 Cm *! 213

Sapindaceae Sapindus saponaria Pod 5-20 Ce *! »* 1

Solanaceae Capsicum annuum - 0.12% Ce ** 198 F po/wh 25-75 Cm *! *! *! 125 F 10-20 Cm *! 218 F 2.5 Cm *i *! 357 - 10 g Cm.Sz ** 22 Solanaceae Capsicum chinense F 10-20 Cm n 218 Solanaceae Capsicum frutescens F 0.03-0.12% Ao, Ce *! *! n 199 F 2g/20seeds Cm ** ** »* ** 59 F po/wh 25-75 Cm ** *! *! 125 F 20 Cm »» ** 132 F - Cm *! *! 160 F 10-20 Cm n *i 218 F 30 Cm, Rd, So, *i 8 Tea, Tg -- 80 ml/1 Cm, Tco ** 354 Solanaceae Capsicum spp. -- 25 Cm *! ** ** 60 Solanaceae Nicotiana tabacum G 25 Cse ** ** ** 67 L 2 g/500 Cm *! *! 209 seeds L 2 Ce, Cm ** *! ** ** *i 245 Solanaceae Solanum incanum -- 50 Cm *i 10

Sphenocleacea Sphenoclea zeylanica Shoot 100-300 Cm *! 93

Tiliaceae Tilia cordata - 1 g/6 beans Ao *! ** ** ** 261

17 2 Powders or fresh application

Plant Family Plant species Plant part Quantity Beetle A O E H L M P Ref.

Verbenaceae Lantana camara -- 5-20 Cc *! *! 50 L po/wh 50 Cc *! 325 L 0.5-4 g/50 Cm ** ** 2 seeds L 10-50 Cm ** ** 69 Verbenaceae Lippia adoensis L 0.5-4 g/50 Cm *! *! 2 seeds Verbenaceae Lippia chevalieri wh Layers Cm ** 59 Verbenaceae Lippia multiflora G 25 Cse ** *! *! 67 Verbenaceae Vitexaltissima L 10-50 Cm *! ** 69 Verbenaceae Vitex negundo Lwh 5-30 Cc *! *i 240 L 20-80 Cc *! *i 267 Verbenaceae Vitexspp . L 10 Ca ** ** 139

Zingiberaceae Aframomum melegueta S 2g/500 Cm ** ** 209 seeds Zingiberaceae Alpinia galanga -- 0.25-3% Cc M H *! 4 R 40 Cm ** 341 Zingiberaceae Curcuma amada -- 0.25-3% Cc M n *! 4 Zingiberaceae Curcuma longa - 1-3% Cc il il n 4 R 10-20 Cc *! *! *! 296 R 20 Cm *! »* 132 Zingiberaceae Curcuma zedoaria -- 0.25-3% Cc M M *! 4 Zingiberaceae Zingiber officinale - 1-3% Cc Ü *! ** 4 R 5 Cm *! *! *! 81 R 20 Cm ** ** 132 R 2g/500 Cm ** ** 209 seeds Zingiberaceae Zingiber spectabile L 100-300 Cc, Cm, Cr *! *! 251 Zingiberaceae Zingiber zerumbet L 100-300 Cc, Cm, Cr *! *! 251

Zygophyllaceae Balanites aegyptiaca R 100-300 Cm *i 93 $: B= bark, Br = root bark, Bs = stem bark, Bu = bulb,F = fruits, Fl = flowers, G = green parts, K= kernels, L = leaves, P = peels, R= rhizomes, roots, S= seeds, T = twigs,Tu = tuber, W = wood; d = among others used dry, fr = fresh, po = among others used as apowder , wh = whole un-ground material used. ~: in gplan t material/kg stored product, unless stated otherwise. A: See table 1 #: A= adult longevity & fecundity, O= oviposition, E = survival of eggs & embryos on the seed surface, H = hatching, L= survival of larvae and pupae inside the seed, M = emergence, P = population numbers & effect on the stored product; ** = Measured, but no statistically significant results were found or presented, *!= Significant decrease, !!= Total inhibition.

18 Theus eo fplan tproduct st oprotec tstore dseed s

2.1 DEVELOPMENTAL STAGE AFFECTED The efficacy ofplan t powders as insecticides depends on theplan t (species,use dparts , harvest time, storage method etc.),th e size ofth epowde r particles (51)an d on the applied quantity.

2.1.1 EFFECTS ON ADULTS Toxicity, either through fumigation orthroug h direct contact, isusuall y the major action ofplan t powders against adult insects in laboratory tests.I n the literature, toxicity levels vary widely, from slight toxicity to induction of complete mortality of all adult insects. Chenopodium ambrosioides and Tephrosia vogeliiaffecte d adult survival of Cse(67) .Leave s andkernel s ofAzadirachta indicaslightl y increased adult mortality of Cm(287) .Fres h and dry whole leaves ofBoscia senegalensis cause d mortality (288) androo t bark ofZanthoxylum zanthoxyloides showed high contact toxicity to adults of Cm(212) . The gum exudate ofAnacardium occidentale mixe d with cowpea flour in artificial seeds reduced the number of surviving adults ofCm (174). Leaves of Pipernigrum caused adult mortality for Cm(248 ) and were highly toxic toSo adults.Th e toxicity was attributable toth epresenc e ofpiperin e (315). Piperguineense seeds caused adult mortality in Cm, Csuan dSz (176). Dennettia tripetalaachieve d complete adult mortality for Sz (215)wherea sAlpinia galanga, Curcuma amadaan d Curcumazedoaria did sofo r Cc(4 )an dRicinus communis for Cm (214). Onth e other hand, repellence accounts for a largepar t ofth e effect ofpowder s of Apiaceae, Lamiaceae and Rutaceae on storagebeetles .Th e effect ofHyptis spicigera, Lippiachevalieri, Ocimum basilicum and Ocimum gratissimum,applie d aslayer s of powderbetwee n layers of cowpea pods,coul db e duet o an insect repellent effect (59). Anethumgraveolens repelle d So, Cman dLs (319). However, in laboratory tests,th erepellen t effect can only be measured ifth etes t insects are given the choicet o escape from thetreate d areas.I n other cases,th etoxi c effects ofplan t powders aremeasured , whereth e effect would berepellen t ifth e insect could get away. Leaves of Ocimum basilicumcause d complete mortality and showed fumigant toxicity against adults ofZs (346),bu twer e found tohav e a repellent effect in other situations(59) . Sometimes both repellent and lethal effects are found. Peels of Citrussinensis an d Citrusparadisi wereal lbot htoxi c andrepellen t to adults ofCm (73 )bu tCitrus sinensiswa smor e effective than Citrusparadisi (9).

2.1.2 EFFECTS ON OVIPOSITION Ifplan t powders reduce adult longevity and fitness, the numbers of eggs laid will often be lower aswell . Moreover, themechanica l effect of large quantities of powders themselves could have aneffec t on oviposition. Inmos t papers,result s are given without anexplanation . Allplan t powdersteste db yJavai dan dMpotokwan e (131) were effective against oviposition of Cm. Capsicumfrutescens, Capsicum annuum and Capsicum chinenseeffectivel y reduced oviposition of Cm(218) ,bu t seeds of Piperguineense (216 ) were more effective than the first two species (125).Th e gum exudate ofAnacardium occidentale mixedwit h cowpea flour in artificial seeds prevented oviposition of Cm(174) .Peel s of Citrusparadisi, Citrusaurantifolia (219 ) and Citruscrematifolia (248) ,roo t bark ofZanthoxylum zanthoxyloides(212) , fruits of Curcuma longaan dEugenia aromatica (132) ,Azadirachta indica, Anacardium occidentalean d Zingiberofficinale all decreased thenumbe r ofegg s laidb y Cm(81) . Mitracarpusscaber, Napoleonaimperialis andDiodia sarmentosa decrease d the

19 2 Powders or fresh application number ofegg s laidb y Cman dSz (176).Leaves ,bar k and seeds ofAphanamixis polystachya deterred oviposition of Cc to some extent (330).Eucalyptus citriodora was effective against oviposition of Cc(325) . Chenopodium ambrosioides and Tephrosia vogeliiaffecte d oviposition ofCse (67) .Leave sofOcimum basilicum suppressed oviposition ofZs completely, but when large quantities ofwhol e intact leaves were applied, oviposition was enhanced (346). Plant powders canhav e an effect on thereproductio n of stored product beetles without affecting the longevity ofparen t orF l adults. Seeds ofAzadirachta indicadi d not impair adult longevity of Cmfemales , but they didreduc e their fecundity (123). Oviposition ofCc an d Cmwa s decreased by leaf dust ofNicotiana tabacum bu t the eggs thatwer e laid developed normally (245).

2.1.3 EFFECTS ON EGGSAN D LARVAE Effects ofplan t powders on eggs are not often found. Curcumalonga exhibite d an ovicidal effect on Cc (4). Larvae of seedbeetles ,protecte d byth etest a and the seed contents,ar eno t very susceptible to the effects ofplan t powders either. Leaves,bar k and seeds of Aphanamixispolystachya decreased larval survival and seed damage by Cc(330) . Piper nigruman d Capsicumfrutescens had an effect on larval development ofAo and Cc(199) . Plant powders often prevent orreduc e the emergence of adult beetles from the seed. However, it isno t clear ifthi s effect iscause d by larval mortality, orb y the fact that the emerging adults contact theplan t powder while gnawing their way out ofth e seed. Allplant s tested by Javaid andMpotokwan e (131)wer e effective against emergence of Cm but none ofthe m decreased the seed weight loss.I nthi s case,th e seed weight loss was said to imply that the larvae did develop completely. Based onth e data provided in mostpublications , it isno tpossibl e tounequivocall y determine the stage ofth ebeetl e that was affected. Anacardiumoccidentale, Zingiberofficinale an d Azadirachta indica(81 ) andpeel s of Citrusparadisi and Citrus aurantifolia significantly decreased infestation and emergence of Cm(219) . Seeds ofAzadirachta indica had the same effects and theyprolonge d the developmental period ofth e beetle aswel l (123).Piper guineense reduce d adult emergence in Cm(216) . Dennettia tripetalaan dPiper guineense inhibited emergence of Cman dSz completely (215). Eucalyptuscitriodora (325 ) and leaves of Vitex negundo(240 )wer e effective against emergence of Cc.See d treatment with wholeplant s of Chamaecrista nigricansan d with some ecotypeso fHyptis spicigera decrease d emergence ofAo (162). In some cases,th eprotectin g effect ofplan tpowder s against insects is mentioned without any indication of the susceptible developmental stage. Whole leaves ofBoscia senegalensiscause d areductio n ofprogen y numbers and adecreas e in seed damage by Cm(288) . Capsicumfrutescens effectively controlled Rd (8).Leave s ofAzadirachta indicaha d antifeedant properties and an inhibiting effect onth e growth and reproduction of storage insects (119). Trigonellafoenum-graecum inhibited larval penetration ofAo and was moderately toxic to larvae of Tea (234). Sometimes plant powders are found tohav e aneffec t opposite ofwha t was aimed for. High concentrations of intact leaves ofOcimum basilicumincrease d hatching and progeny emergence (346).

20 Theus eo fplan tproduct st oprotec t storedseed s

2.2 COMPARISON OF BEETLE SPECIES Not all storagepes t insects areequall y susceptible toth eeffec t ofplan tpowders . Capsicumfrutescens andHyptis spicigera effectivel y controlledRd but wereno t effective against Cm,So, Tea and Tg(8) . Citrussinensis an d Citrusparadisi peels were more toxic to Cmtha n toDm adults (73).Mitracarpus scaber, Napoleona imperialisan dDiodia sarmentosa prevente d oviposition of Cman dSz bu t not ofCsu (176). Thetes t conditions can influence theoutcom e ofexperiments . In laboratory tests,Ao was most effectively controlled by an oil extract ofLavandula angustifolia an d for the control ofSg, drie d dust ofLaurus nobilis wa s most effective. However, under storehouse conditions,th e best insecticidal efficacy against Sg inmillin g wheat was shown by dust ofRosmarinus officinalis, wherea s in seed wheat an oil extract of Laurusnobilis wa sbes t(138) .

2.3 COMPARISON OFPLAN T SPECIES Results oftest swit hneem ,Azadirachta indicaar e sometimes ambiguous. Usually neem isver y effective against insects.Th e seeds were more insecticidally effective than Citruschinensis an dEupatorium odoratum (12).Bu tno t always isAzadirachta indicaamon g the most effective plants.Azadirachta indica, Zingiberofficinale an d Curcuma longawer e lesseffectiv e thanAlpinia galanga, Curcuma amadaan d Curcumazedoaria against Cc(4) .Anacardium occidentale protecte d cowpea better against Cmtha nZingiber officinale and/'orAzadirachta indica di d (81). Moreover, some contradicting results have alsobee n found inth e screened literature. According to Chiranjeevi and Sudhakar (50),Azadirachta indica completel y inhibited the development of Ccwherea sAcorus calamus di d so only slightly. However, Azadirachta indicawa s less effective against Cctha nAcorus calamus kerne l (258) and lessrepellen t than twigs ofAcorus calamusan dLedum palustre (121).Acorus calamus,Thevetia peruviana andIpomoea carnea effectivel y protected seeds against Cc(228) .Azadirachta indicaan dZanthoxylum zanthoxyloideswer e both as effective aspirimiphos-methy l andpermefhri n inreducin g bruchid damage (211),bu t Ogunwolu andOdunlam i (212)foun d Zanthoxylum zanthoxyloidest ob e significantly more effective against Cmtha nAzadirachta indica. Pepper species, Capsicuman dPiper, ar etraditionall y often used. When comparedt o other plants they are often, but not alwayshighl y effective. Here again, contradictory resultshav ebee n found. Ofsi xplants , Capsicum annuum wasth e onlyon etha t effectively controlled Cman dSz (22). Capsicum annuum,Capsicum chinense an d Capsicumfrutescens inparticula r were effective inreducin g damaget o cowpeab y Cm(218) .Th e insecticidal properties of Capsicumfrutescens werebette r than carbon- bisulphite andpyrethru m (339).However , according toZehre r (354), Capsicum frutescens was ineffective against Cman d Tcoan dth e results did not differ from the untreated control.Besides ,Morallo-Rejesu s etal. (199) found thatPiper nigrum showed more contact toxicity againstAo and Cctha n Capsicumfrutescens. Piper nigrumreduce d damage by Cm,wherea s Capsicumfrutescens, Zingiberofficinale an d Alliumsativum wer e not effective (132).Piper guineense enhance d mortality inCm (178) more than Capsicumannuum an d Capsicumfrutescens (125) and it caused mortality in Csuan d Sz adults (176). However,Dennettia tripetala wa s more effective than Piperguineense, Monodoramyristica an dXylopia aethiopicaagains t Sz(215) . Forplant s ofth e Lamiaceae,th eresult s oflaborator y testsvar y greatly. DryHyptis spicigeraan d Chamaecrista nigricansdi dno t decrease oviposition ofAo, but some

21 2Powder so rfresh applicatio n

ecotypes did decrease emergence (162).Thes epowder s reduced infestation byCm considerably (339) andHyptis spicigera effectivel y controlledRd but itwa s not effective against other seedbeetles .Hyptis suaveolens wa s not effective against anyo f the insect speciestested , including Cm(8) ,bu t often plants tested,Hyptis suaveolens andSphenoclea zeylanica showed the bestprotectan t effects against Cm(93) . Ocimum basilicuman dLippia multiflora, Eupatoriumodoratum an dNicotiana tabacumha d little orn o effect on Cse(67) .Rosmarinus officinalis, Origanum vulgare, Thymus vulgarisan dLaurus nobilis wer e only slightly effective against Ao butunde r storehouse conditions,Rosmarinus officinalisshowe d the highest insecticidal efficacy against Sg (138).Mentha spicata wa sa seffectiv e astw o chemical pesticides against Ca(100) .O fMentha piperata, Rosmarinusofficinalis, Thymus serpyllum, Thymus vulgaris,Allium sativum, Cymbopogon nardus, Eucalyptusglobulus, Laurus nobilis an dSatureja hortensis, the first five, all Lamiaceae,provide d thebes t insecticidal effect againstAo and among them,Thymus serpyllumwa sth emos t efficient (261). Plantsfrom othe r families can also showdifferin g rateso fefficacy . Thevetia peruviana andIpomoea carnea effectivel y protected seeds from damageb y Cc, whereasAdhatoda vascialeave sprove d tob e ineffective (228).Eugenia aromatica reduced damageb y Cmcompletel y whereas Curcuma longareduce d only oviposition (132).Leave s ofEugenia unifloraan dLippia adoensis wer e effective against Cm,bu t Lantanacamara an d Cymbopogon citratuswer eno t (2).Dr y and fresh leaves,an d wholean d ground seeds ofSenna occidentalis did not show contact toxicity toCm and didno tprotec t stored seeds (165).Non e ofth eplan t powders tested byJavai d and Mpotokwane (131) decreased the seed weight loss duet o Cminfestation . Annona senegalensis, Entadaafricana, Khayasenegalensis an dParkia biglobosawer e ineffective oreve n increased thenumbe r ofdamage d seeds(93) .Al l plant powders tested byJavai d and Mpotokwane (131)wer e effective against Cm,bu tTerminalia sericeaan dPeltophorum africanum wereth e only ones inhibiting adult emergence. Citrussinensis wa smor eeffectiv e than Citrusparadisi asa repellen t anda toxicant against adults of Cm(9 ;73) . Theplan t part and itspreparatio n used inexperiment s can influence theresults . Seeds ofAphanamixis polystachya weremor eeffectiv e against Cctha n the leavesan dbar k ofthi splan t (330).Fres h ground leaves ofBoscia senegalensis wer e more effective than fresh entire leaves or dry leafpowde r against Cm(288) .A t low dosages, kernels ofAzadirachta indicawer e more effective against Cm than leaves (287).Bruchi d mortality washighes t for ground fruits of Capsicum annuum,lowe r for sliced fruits and lowest for whole fruits (357).

2.4DURATIO N OFTH E EFFECTS Thepersistenc e ofpowder s orfres h plants isusuall y better than for instance that of volatileoil s(chapte r 4).I nmos t studies,th eshort-ter m effect is lessimportan t than theoveral l long-term effect onth ebeetl e population. Ricinuscommunis leave s (214), Capsicum annuum(22 ) andDennettia tripetala, Piperguineense, Monodora myristicaan dXylopia aethiopica(215 )offere d seedprotectio n for 3months .Hyptis suaveolensan dSphenoclea zeylanica stil l showed protectant effects after 4 months (93). Grain treatment withPiper nigrum seed s caused areductio n damage byAo after 4 months (163). Thevetiaperuviana, Acoruscalamus an dIpomoea carnea effectively protected seedsfrom damag eb y Cc,fo r atleas t 135day s (228).Th e effect of Zanthoxylum zanthoxyloidesan dAzadirachta indica agains t Cmlaste d for nearly 5

22 The use of plant products toprotec t stored seeds

months (212).Residua l toxicity ofPiper nigrum and Capsicumfrutescens lasted for 6 months (199)an d Capsicum annuum effectively controlled Sz upt oth e sixth month (22).Kernel s ofAzadirachta indica protected cowpea against Cmsatisfactoril y up until 8month s (311). Vitex negundo reduced oviposition and adult emergence ofCc up until 9month s (240).Azadirachta indica kernel powderprotecte d stored seeds for up to 11month s against Cm(134) .Leave s of Cissus quadrangularisprotecte d cowpea from insect pests for 1-2 years,wherea sSwartzia madagascariensis did so for 1year ,Prosopis africana fo r 3-5 years andPterocarpus santalinus an dMaerua angolensisfo r up to 7years ,an d the latter did not show any effect on germination (153).

2.5 EFFECTS ON THE STORED PRODUCT Powders mayhav e an effect onth e seedsthe y are supposed toprotect . Seeds treated withMonodora myristica change d colour (215)an dpowdere d fruits ofCapsicum annuumlef t are d colour onth e seeds (357). Stored grain,treate d withAzadirachta indicasee dpowder , was spoiled duet o growth ofth enee m seed-borne fungus ofth e Aspergillus family (236). However, powders do not usually have these adverse effects. Inmos t experiments, the seedsretai n their viability and culinary properties.N o adverse effect on seed germination was found after treatment with Origanum vulgare, Laurusnobilis, Thymus vulgaris, Rosmarinusofficinalis (138) , Thevetiaperuviana,Adhatoda vascia, Acorus calamus, Ipomoea carnea(228 ) andMaerua angolensis (153) . Seed quality and viability wereno t affected by treatment with peels of Citrusparadisi andCitrus aurantifolia(219 ) orb y Capsicum annuum,Capsicum chinensean dCapsicum frutescens (218).Neithe r ofth e treatments with Piperguineense, Capsicum annuum or Capsicumfrutescens affected seed viability or cooking properties (125). Seed germination and taste were preserved quitewel l after treatment with rootbar k of Zanthoxylumzanthoxyloides (212 )an dthes epropertie s were not impaired by treatment withAzadirachta indica kernel powder either (134). Seedsremaine d viable and their texture, colour and overall attractiveness remained unaffected after treatment with seeds ofAzadirachta indica (123).

2.6 CONCLUSIONS Forpowders ,hardl y any general rule ist ob e found. There isn orankin g order for efficacy of theplant s and the best mode ofapplicatio n isunknown . The preparation and application are easy andtherefor e manyplant s have beenuse d aspowder s inpilo t tests.Applie d quantities and results from tests vary greatly. Ifth eright plan t is chosen, all stages ofth e developing beetle canb e affected. If aneffec t is found, it usually lasts for a few months. There areusuall y no effects on the stored product.

23 2Powder so rfresh applicatio n

24 Theus eo fplan tproduct st oprotec t storedseed s

3 ASH Whereas the effect ofpowder s ismainl y toxic orrepellent , the effect of ashmixe d with the stored seeds,a traditional means ofprotection , ismainl y physical. Ashes of different plant species can have different effects on insects duet o their structure andparticl e size or to components of theas h that still havetoxi c effects onth ebeetles .Generally ,th e finerth eparticle s are and/or thegreate r the applied quantity, themor e effective this method will be.Th e quantities of ashneede d to effectively protect stored products are large.Th e treatment will occupy much ofth e storage room andwoul d therefore be useful for small quantities ofcowpe a only(59) .Th eplant suse d asashe san dthei r effects on storage beetles are summarised intabl e3 .

Table3: Incinerated material used against storage insects, the quantity andits effect. Plant (sub-) Plant species Quantity ~ Beetle species A Affected stage # Refe­ family A O E H L M P rence

Bombacaceae Ceiba pentandra 1000 ml/1 Cm *! *i 349

Casuarinaceae Casuarina indica 10-20 Cc *! *! 50

Combretaceae Combretum imberbe 60 Cm *! *! *! 131 20 Cm *! *! 132 100-1000 Cm *! *! *! 133

Leguminosae- Afaelia africana 2g/20 grains Cm ** *! ** ** 59 Caesalpinioideae 2:3-1:1 v:v Cm *i *! ** 60 1000 ml/1 Cm *! *! 349 Leguminosae- Tamarindus indica 20-25 Cm, Rd, So, Tea, Tg *i 8 Caesalpinioideae

Leguminosae- Acacia nilotica 10-20 Cc *! *! 50 Mimosoideae Leguminosae- Acacia spp. 50 Cm *i 10 Mimosoideae Leguminosae- Parkia biglobosa 1000mV l Cm *! *i 349 Mimosoideae

Leguminosae- Phaseolus spp. 50-300 Cm *! *! *! 140 Papilionoideae Leguminosae- Pterocarpus erinaceus 2g/20 grains Cm ** *! ** ** 59 Papilionoideae 2:3-1:1 v:v Cm *i *! ** 60

Meliaceae Azadirachta indica 10-20 Cc *! *! 50 20 Cm,Sz •* 22

Poaceae Cymbopogon nardus 5-100 Ca *! *! 139 Poaceae Oryza saliva 40 Cm *! 247

25 3 Ash

Plant family Plant species Quantity Beetle A O E H L M P Ref. Poaceae Pennisetum spp. 2g/20 grains Cm ** *! ** ** 59 2:3-1:1 v:v Cm *! *! ** 60 Poaceae Sorghum bicolor 1000 ml/1 Cm *! *! 349

Rubiaceae Coffea arabica 33-66 Ao *! 338

Sapotaceae Vitellaria paradoxa 2g/20 grains Cm ** *! ** ** 59 2:3-1:1 v:v Cm *! *! ** 60 1000 mV I Cm *! *! 349

Cattle dung 10-20 Cc n *! 50 25-1000 Cm *! *! *! 133

Wood (kitchen stove) 200 Ao *; 30 33-66 Ao *! 338 0.25-1.001/1 Ao, Zs *! *! 49 50 Ao, Zs ** ** 309 10% Ca ** *! 100 2g/20 grains Cm ** *! ** ** 59 500 ml/1 Cm *! 66 10-40 Cm n M 341 1000 ml/1 Cm *! *! 349 0.125-1.01/1, Cm n ** 350 mix & layer 20 Cm.Sz ** 22 ~:Quantit y ing ash/k gstore dproduct ,unles sstate dotherwis e A:Se etabl e 1 #:A =adul tlongevit y& fecundity, O =oviposition ,E = surviva lo fegg s& embryoso nth esee d surface, H= hatching ,L= surviva lo flarva ean dpupa einsid eth eseed ,M =emergence ,P = populatio nnumber s& effect onth estore dproduct ;* *= Measured ,bu tn ostatisticall y significant resultswer efoun d or presented, *! =Significan t decrease, !! =Tota linhibition .

3.1 DEVELOPMENTAL STAGE AFFECTED

3.1.1 EFFECTS ON ADULTS Most ofth e effect ofashe s iscause d by amechanica l rather than by achemica l action. Immobilisation ofth e adult insectsplay s an important role.Adul t beetles use the inter- granular spacet o infest the stored product. Obstruction of inter-granular spaces hinders movement ofth e adult beetles and thus leads to less or shallower infestation (110). Sinceth emovemen t ofth e adults ishampere d (66; 140)thei r latitude ofmovemen t and ofmeetin g conspecifics is limited (339) andthei r rate ofmultiplicatio n will thusb e lower.Th eevolutio n ofth e infestation inash-treate d stored seeds isslowe rtha n in untreated control samples.Th e bruchids go toth e surface when they can and lay their eggs on seeds sticking out of theas h (60).Accordingly , any dry,powder y substance, such asfin e sand (66), filling the inter-granular space might serve as agoo d protective medium for stored seeds.However , the relatively heavy sand particles, and powders of

26 Theus eo fplan tproduct st oprotec t storedseed s irregular sizecanno t be evenly mixed with thebean s and aretherefor e less effective than fine ash(49) . The applied ash doesno t onlyhampe r beetle movement, but itca n alsod o physical damage to the adult beetles.I fth e adult insectsmov e over orthroug h the ash,thei r bodies (22;65) ,especiall y the layer ofchiti n onth e adults' abdomen are grazed (339). Clogging of insect spiracles andtrachea e (350) orblockin g ofth e lateral stigmates,al l essential for respiration, cause suffocation ofth e adult and enhance mortality (66). Toxicity and/orrepellenc e by (components of) the ashes could alsob e important inth e effects against insects.I n this case,i t doesmatte r which material is incinerated to use against bruchid infestation. Remains oftoxi c compounds in ashes of specially selected plants might have extra effects onth ebeetles . Millet ash was saidt o be effective, because it contains acidic components that repel thebeetles .Farmer s recommended ashes ofPterocarpus erinaceus, Vitellariaparadoxa, Pennisetumspp .an dAfzelia africanafo r their specific essence (59;60) . Katanga Apuuli and Villet (140) found that theus e ofmois t materials could induce premature germination ormoul d of thestore d seeds.Thi s is,however , inconsistent with the finding that ash canprotec t stored seeds from mould (153),an d with the experience that the layer of ash covering the contents of storagestructure s is sometimes drenched with water to close itbette r andt omak e itmor e airtight (66).

3.1.2 EFFECTS ON OVIPOSITION If applied in large enough quantities,th e effect of asho n oviposition can be important. Wood ash failed topreven t oviposition of Ca (100),bu t Cmlai d fewer eggs on beans treated with ash (350) of Combretum imberbe (131; 132)o r of cow dung than on untreated control beans (133).Decrease d oviposition could be caused by the restriction of movement ofth ebeetle s among the seeds,thu s diminishing thepossibilit y to oviposit directly onto the seed (140).As h was alsoreporte d to obstruct the adhesion ofth eeg gt o the grain,jus t asnon-volatil e oils did (119),bu t the effect on oviposition could alsojus t be attributed toth e shorter lifetime ofth e females (59).

3.1.3 EFFECTS ON EGGSAN D LARVAE Suffocation could be amechanis m ofth e effect of asho n eggs,adult s and larvae. Thick layers ofas h reduce the available oxygen (339) and could interfere with the respiratory ability of eggs, larvae and adult bruchids (140). However, themechanis m ofth e effect ofas h isno t always clearly stated. Ash ofco w dung (133) and of Combretum imberbesignificantl y reduced adult emergence ofCm (131), asdi d ordinary wood ash (208).Larva l andpupa l mortality of Cmwer ehighe r in beans treated with ash than inuntreate d controls (350).As h was more effective than sand and millet husks preventing adult emergence ofZs andAo (49).

3.2 COMPARISON OFBEETL E SPECIES Since the effect of asheso n storage beetles ismostl y mechanical, the differences in susceptibility between different beetle species depend mostly on thedifference s inthei r life cycle.Ba leavesth e store after a few generations,wherea s Cmstay s inside (119). Therefore, it couldb etha t the latter species ismor eheavil y affected bytreatment s with ash.Sz was more affected andthu s caused less damage to seeds treated with ashtha n Cm(22) . Ao was more affected by ash than Zs(49) . The differences in efficacy can depend onth eproduc t that istreate d rathertha n onth e differences in susceptibility between beetle species.Azadirachta indicaas h and ordinary

27 3 Ash

wood ash did not control Cm.Th e smooth surface ofth ecowpe a prevented adherence of the applied ash toth e seed, whereas onth eroughe r seed-surface of maize,use d for tests withSz, th e ash adhered better (22).

3.3 COMPARISON OFPLAN T SPECIES Significant differences inweigh t loss and innumber s ofbeetle s were noted for different ashes.Parkia biglobosaan dAfzelia africana were more effective thanVitellaria paradoxa, Ceibapentandra, Sorghumvulgare, o rmixture s of theseashes . Chemical analyses ofthes e ashes couldno tprov e thepresenc e ofheav y metals or alkaline saltsa s insecticidal components.However , tests with sand ofth e sameparticl e size asth e ashes did not give similarresults , soth emer epresenc e of inertparticle s ofdifferen t sizes couldno tb eth e (only) cause for the differences inth e results (349).Co w dung ash was more effective than Combretum imberbeas h (133).However , Javaid & Mpotokwane (131) found thatth e ash of Combretum imberbe could easily and maybeeffectivel y be used ifi twer e integrated with insecticidal plantproduct s orwit hresistan t seeds. Theresult s are often given without explanations ofth epossibl e mechanisms of action. Ash ofPhaseolus spp.harvest-remain s protected cowpea.A mixtur e ofas h and beans gave very satisfactory protection, with fewer offspring than there were parents(140) . Wood ash was found to be more effective than onion scale leaves and dry chilli pepper fruits (208).

3.4 DURATION OFTH E EFFECTS Ash seems tob e appropriate for storage periods of intermediate lengths.A to p (a middle) and abotto m layer will be enough topreven t (new) infestation. Wolfson etal. (350) suggested a 1:1 (by volume) ratio ofas h with alaye r on topt oprotec t the cowpeas for at least onegeneratio n ofbeetles .Azadirachta indicawoo d ash and ordinary wood ash controlled Szu p to five and sixmonth s respectively (22).As h would be appropriate for athre e to sixmonths ' storage period (208) and kitchen ash caused bruchid damage toremai n belowth e economic threshold for at least 9month s (30).Thi s isi n contrast withfindings from anothe r study inwhich , after four months of storage,th e application ofwoo d ash did notresul t infewe r beans tob e damaged and there were not significantly fewer beetles than inth euntreate d control experiment (309).

3.5 EFFECTS ON THE STORED PRODUCT There are a few clear disadvantages associated withth eus e ofas h for protection of stored products. Duet oth e ashtreatment , the appearance ofth e beans can change.As h ofSorghum vulgare change d the colour ofth ebeans ,wherea s other ashes did not do so (349). Farmers havereporte d that wood ash affected the marketability of the stored product (309)becaus e itmad e thebean s appear old and dirty.As h could alsohav ea negative effect on germination ofth e stored seeds (31), although George and Patel (100) found that thepercentag e ofgerminatio n of green gramtreate d with ash was higher than for the untreated control.Anyhow , the applied ash should bewel l dried to prevent the induction ofprematur e germination or moulding,bu t then, due to the dehydrating action ofth e ash,bean s can become very hard and therefore incookable and inedible (140).

28 Theus eo fplan tproduct st oprotec tstore dseed s

3.6 CONCLUSION Ash can provide an effective, simple, cheap and clean way ofprotectio n of stored seeds against storage beetles, if it isapplie d in large enough quantities. Care shouldb e taken that both thebean s and the ash are very dry from the beginning ofth e storage on,t o prevent moulding. For small quantities ofbeans , thismetho d could beuseful , butth e quantities of ash and storage space,neede d toprotec t large quantities of cowpea, areno t easily available.

29 Theus eo fplan tproduct st oprotec t storedseed s

4 VOLATILE OILS From some aromatic plants volatile oils canb e extracted. These oils canb e applied to stored seeds asprotectant s against storage insect pests.Th eyiel d ofoi l isusuall y low, but duet orepellenc e ortoxicity , even small amounts ofth econcentrate d essential extract can be very effective inairtigh t orhermeti c storage structures.A major advantage ofvolatil e insecticides istha tthe y dono t need tob emixe d with theseeds . Nophysica l contact isneede d between seedsan dprotectant . Theeffec t ofthes e volatile oils isusuall y reached through fumigation. All plants discussed inthi s chapter and in table 4 areuse d asvolatil e oils,unles s stated otherwise.

Table 4:Plants ofwhich the volatile oil isused against storage insects,the quantity of oil usedand its effect. Plant (sub-) Plant species Quantity ~ Beetle species A Affected stage # Refe­ family A O E H L M P rence

Annonaceae Dennettia tripetala 3 ml/kg Cm, Sz H ** 215 Annonaceae Monodora myristica 3 ml/kg Cm.Sz ** ** 215 Annonaceae Xylopia aethiopica 3 ml/kg Cm.Sz ** ** 215

Apiaceae Anethum graveolens 5-10 g/kg Ao ** n 262 Apiaceae Apium graveolens 5-10 g/kg Ao *! n 262 Apiaceae Coriandrum sativum 5-10 g/kg Ao *! *! 262 10-50 ug/insect Cm, Ls, So, Tco *i 320 Apiaceae Cuminum cyminum 5-10 g/kg Ao *! n 262 Apiaceae Diplolophium africanum 6.7-33.3 Cm *! *! n 106 6.7-33.3 Cm ** *! 143 44.8 mg/1 Cm, Csu !! n 154 Apiaceae Petroselinum crispum 5-10 g/kg Ao *! n 262

Araceae Acorus calamus 2-4% Cc *! 111 1-7 ul/insect Cc *! 332 10-50 ul Cc, Rd, Sg, So, Tco *! 87 2.5-125 Cc, Rd, Sg, So, Tco *! *! 278 io ni Cc, Sg, So *! *! 280 10-50 ni Cc, Sg, So, Tco *! ** ** 264 25-125 Cc, Sg, So, Tco *! *! 279 12.5-25 Cph It *! *! *! 246

Asteraceae Ageratum conyzoides 0.1-2 g/kg Ao, Cc n H 199 Asteraceae Blumea balsamifera 0.1-2 g/kg Ao, Cc M il 199 Asteraceae Chromolaena odorata 0.5-30 ul/50 seeds Cm ** ** 2 5-20 nl/50 seeds Cm *! *! 99 Asteraceae Chrysanthemum indicum 0.1-2 g/kg Ao, Cc Ü n 199 Asteraceae Eupatorium capillifolium 20-120 Cm *! 25

31 4Volatil eoil s

Plant family Plant species Quantity Beetle A O E H L M P Ref. Asteraceae Tagetes minuta 4-37.5 ul/insect Cm *i *i il il 141

Geraniaceae Pelargonium spp. 1-10% dip Cc *! *! !! n 263

Lamiaceae Coleus amboinicus 0.1-2 g/kg Ao, Cc u n 199 Latniaceae Hyptis spicigera 0.2-0.35 ul/insect Cm *! 72 Lamiaceae Hyptis suaveolens 0.7-2.2 ul/insect Cm *! 72 4-37.5 ul/insect Cm ** *! H !! 141 Lamiaceae Lavandula angustifolia 0.1-0.74 ml/kg Ao.Sg *! 138 5-10 g/kg Ao ** n 262 Lamiaceae Lavandula spp. 6.7-33.3 Cm *! *i n 106 6.6-33.3 Cm il M 142 Lamiaceae Mentha citrata 5.5-174 Cc *i n 185 Lamiaceae Mentha piperita 5.5-174 Cc *! n 185 5-10 g/kg Ao ** ** 262 Lamiaceae Mentha spicata 5.5-174 Cc *! H 185 Lamiaceae Ocimum basilicum 5-10 g/kg Ao ** n 262 1-10% dip Cc *i *! n M 263 0.05-0.17ul/insect Cm *! 72 1-37.5 ul/insect Cm *! *! ** *! 141 0.2-2 ml/kg Cm, So *! 96 Lamiaceae Origanum majorana 5-10 g/kg Ao ** n 262 Lamiaceae Origanum vulgare 5-10 g/kg Ao *! n 262 0.1-0.74 ml/kg Ao.Sg *! 138 Lamiaceae Rosmarinus officinalis 5-10 g/kg Ao *! n 262 0.1-0.74 ml/kg Ao.Sg *i 138 Lamiaceae Salvia officinalis 5-10 g/kg Ao ** n 262 Lamiaceae Satureja hortensis 5-10 g/kg Ao *! u 262 Lamiaceae Thymus serpyllum 5-10 g/kg Ao *! n 262 Lamiaceae Thymus vulgaris 5-10 g/kg Ao ** H 262 0.1-0.74 ml/kg Ao.Sg *! 138

Lauraceae Cinnamomum mercadoi 10-50 mg Cc *! 98 Lauraceae Cinnamomum verum 5-10 g/kg Ao *! n 262 Lauraceae Laurus nobilis 5-10 g/kg Ao ** ** 262 0.1-0.74 ml/kg Ao.Sg *! 138 Leguminosae- Galega spp. 1-10% dip Cc *! *! *! *! 263 Papilionoideae

32 Theus eo fplan tproduct st oprotec tstore dseed s

Plant family Plant species Quantity Beetle A O £ H L M P Ref.

Myristicaceae Myristica fragrans 5-10 g/kg Ao ** M 262

Myrtaceae Callistemon citrinus 20-120 Cm *! 25 Myrtaceae Eucalyptus citriodora 6.7-33.3 Cm *! *! *! 106 6.6-33.3 Cm !!! M 142 6.7-33.3 Cm ** *! 143 0.2-2 ml/kg' Cm, So M 96 Myrtaceae Eucalyptus globulus 5-10 g/kg Ao ** ** 262 Myrtaceae Eucalyptus spp. 1-10% dip Cc ** *! *! *! 263 Myrtaceae Eucalyptus tereticornis 0.2-2 ml/kg Cm, So »* 96 Myrtaceae Eugenia aromatica 0.75-1% w:w Cc u 111 1-3 Cm ** ** ** *• 129 12.5-100 mg/kg Cm *! ** ** 158 Myrtaceae Eugenia uniflora 0.5-30 nl/50 seeds Cm *! *! 2 5-50 ul/50 seeds Cm *! *! 99

Pinaceae Cedrus deodara 1-3% Cc M ** 244 Pinaceae Cedrus spp. 6.7-33.3 Cm *! *! *! 106 Pinaceae Pinus spp. 3% Cc.Sz *! 200

Piperaceae Piper acutifolium 66.7-133.3 Cm ** *! 166 Piperaceae Piper guineense 2-3 Cm M *! M 124 4-37.5 ul/insect Cm »! *! *! n 141 0.002-0.8% Cm M M 216 3 ml/kg Cm.Sz n ** 215 Piperaceae Piper nigrum 0.2-0.8% Cm *! 1! 249

Poaceae Cymbopogon citratus 1-10% dip Cc *! *! *! *! 263 0.5-30 ul/50 seeds Cm Ü H 2 2-8 g/kg Cm *! 36 5-40 ul/50 seeds Cm Ü *i 99 6.7-33.3 Cm *! *! M 106 6.6-33.3 Cm M M 142 6.7-33.3 Cm ** *! 143 4-8 g/kg Cr *! 250

33 4Volatil eoil s

Plant family Plant species Quantity Beetle A O E H L M P Ref. Poaceae Cymbopogon nardus 5-10 g/kg Ao ** M 262 1-10% dip Cc *! *! *! *! 263 2-8 g/kg Cm *! 36 6.7-33.3 Cm *! *! H 106 6.6-33.3 Cm M M 142 6.7-33.3 Cm *i *! 143 25-200 mg/kg Cm *! ** ** 158 Poaceae Cymbopogon 6.7-33.3 Cm *i *! u 106 schoenanthus 6.6-33.3 Cm M !! 142 6.7-33.3 Cm *! *! 143

Rutaceae Citrus aurantifolia 1.6-50 ng/insect Am, Cm, Ls, So, M 324 Tea, Tco 2.5-10g/k g Cm *! *! n 323 3.5-14 ml/kg Cm, Dm, Sz *! ij H 77 2-20 Cm, Dm, Sz *! 78 4-40 Cm, Dm, Sz *i *! 79 Rutaceae Citrus limon 1.6-50 ng/insect Am, Cm, Ls, So, ii 324 Tea, Tco 5-10 g/kg Ao ** *i 262 1-10% dip Cc ** *! ** *! 263 0.1125-0.9g/k g Cm *i ** ** 158 50 g/1, 0.5 nl/insect Cm M 314 2.5-10g/k g Cm H !! n 323 4-40 Cm, Dm, Sz *i *i 79 Rutaceae Citrus paradisi 1.6-50 ng/insect Am, Cm, Ls, So, H 324 Tea, Tco 2.5-10g/k g Cm *! ij n 89 2.5-10g/k g Cm *! *! M 323 3.5-14 ml/kg Cm, Dm, Sz *! *! *! 77 4-40 Cm, Dm, Sz *! *! 79 Rutaceae Citrus reticulata 1.6-50u.g/insec t Am, Cm, Ls, So, ** 324 Tea, Tco 2.5-10g/k g Cm *! 1! n 323 3.5-14 ml/kg Cm, Dm, Sz *! *! *! 77 4-40 Cm, Dm, Sz *! *! 79

34 Theus eo fplan tproduct st oprotec tstore dseed s

Plant family Plant species Quantity Beetle A O E H L M P Ref. Rutaceae Citrus sinensis 1.6-50 ug/insect Am, Cm, Ls, So, ** 324 Tea, Tco 2.5-10g/k g Cm *t *! *! 89 2.5-10g/k g Cm *! M M 323 4-40 Cm, Dm, Sz *! *! 79 0.2-2 ml/kg Cm, So ** 96 Rutaceae CitrusX tangelo 1.6-50 ug/insect Am, Cm, Ls,So, ** 324 Tea, Tco 2.5-10g/k g Cm *! *! *! 323 Rutaceae Fortunella spp. 1.6-50 ug/insect Am, Cm, Ls, So, *! 324 Tea, Tco 2.5-10g/k g Cm *! !! M 323 Rutaceae Murraya koenigii 3.4-3400pp m Cc M Ü *! 232

Verbenaceae Lantana camara 0.5-30 ul/50 seeds Cm *! *! 2 5-40 ul/50 seeds Cm *! ** 99 Verbenaceae Lippia adoensis 0.5-30 ul/50 seeds Cm Ü H 2 5-15 ul/50 seeds Cm M M 99 Verbenaceae Lippia multiflora 6.7-33.3 Cm *! *t M 106 6.6-33.3 Cm M M 142 6.7-33.3 Cm ** »! 143 6.28-12.56 rag/1 Cm *! M M 155 Verbenaceae Verbena officinalis 5-10 g/kg Ao »* *! 262 Verbenaceae Vitex negundo 0.1-2 g/kg Ao.Cc M M 199 ~:Quantitie si nn ioil/ 1 ofai runles sstate dotherwis e A:Se etabl e 1 #:A =adul tlongevit y& fecundity , O =oviposition ,E = surviva lo fegg s& embryo so nth esee dsurface , H= hatching ,L =surviva lo flarva ean dpupa einsid eth eseed ,M = emergence ,P = populatio nnumber s & effect onth estore dproduct ;* *= Measured ,bu tn ostatisticall y significant resultswer efoun do r presented,* ! =Significan t decrease, !! =Tota linhibition .

4.1 DEVELOPMENTAL STAGE AFFECTED

4.1.1 EFFECTS ONADULT S Volatile oilsmostl y affect adultbeetles .Th e vapours usually have arepellen t effect, causingth ebeetle st ofle e from the store,o rno tt oinvad e ita tall .I nmos t laboratory set-ups,however , thebeetle s cannot escape from thetes t arena. This couldb ea caus e for the frequent findings oftoxi c action ofvolatil e oilsagains t adultbeetles . Formulations ofAcorus calamusal lshowe d anexcellen t knockdown effect onCc an d a long lastingresidua l effect (351).Dennettia tripetala an dPiper guineense achieved complete adult mortality forCm an dSz (215) .Lippia multiflora wa stoxi c toadult so f On (155).

35 4Volatil eoil s

Fumigation often leadst orepellenc e ortoxicity . As fumigants inrelativel y enclosed or airtight systems, essential citrus oils couldb e efficient to control insects.Thei r contact toxicity isno t important for the effect (77).Adult s of Cc were susceptible to vapours of Acorus calamus(278) .I n fumigation experiments, Cymbopogon schoenanthusan d Lavandula spp.wer e very toxic to Cmadult s(106) . However, contact toxicity hasbee n found to alsopla y an important role against beetles. Topical application ofeigh t lyophilised citruspee l oils showed high toxicity toCm adults (324).Ageratum conyzoides, Blumeabalsamifera, Chrysanthemum indicum, Coleus amboinicusan d Vitex negundoexhibite d contact toxicity toth e same beetle species,an d the oilswer e even more effective when mixed with the seeds (199). Significant mortality of Cmadult s occurred in fumigation bioassays withCallistemon citrinusan dEupatorium capillifolium(25) .Fro m lemon peel oil three fractions were isolated thatwer emor etoxi c to Cmadult s than the crude oil,upo n topical application (314).

4.1.2 EFFECTSO N OVIPOSITION The number ofegg s laid is often reduced after treatment with volatile oils.Thi s effect on oviposition can becaused , among others,b yth ereduce d longevity ofth e adult insects. Citruspeel s reduced oviposition through adult mortality, but hadn o residual activity onth e eggs or larvaeproduce d by surviving adults (77). Ocimum basilicuman d Pelargoniumspp .cause d areductio n ofovipositio n and of adult longevity (263). However, the effect isno t always due to areductio n ofadul t longevity. For Cm,Piper acutifoliumha d no effect on adult mortality, but it did cause a decrease in oviposition (166).Th e effect ofoil s on female fecundity has been studied in detail.Fo r Cc, Acorus calamusinduce d infertility andregressio n in the terminal follicles ofth e vitellarium initially, and later a shift to regression of theuppe r parts ofth e ovary. Derailment of follicular epithelium was also observed, which resulted into the irreversible atrophy of the follicles (332). In most cases,a reductio n of oviposition ismentioned , without thepossibl e cause. Cymbopogon citratus,Eugenia uniflora, Lantanacamara an dLippia adoensis completely inhibited oviposition and adult emergence of Cm(2) .Piper guineense (216) , Cymbopogon citratus,Cymbopogon nardus(36 ) andLippia adoensis completel y inhibited oviposition of Cmwherea sEugenia uniflora, Lantana camaraan d Eupatoriumodoratum di d sopartl y (99). Seed treatment withAgeratum conyzoides, Blumeabalsamifera, Chrysanthemum indicum,Coleus amboinicus an d Vitex negundo inhibited oviposition (199).Diplolophium africanum prevented oviposition by Cman d Csu(154) .Piper guineense strongly reduced oviposition, and completely prevented reproduction ofCm (124) . Enhancing effects ofvolatil e oils on oviposition have alsobee n found: median- andsub ­ lethal doses ofEugenia aromatica, Cymbopogon nardusan d Citruslimon cause d oviposition tob e enhanced for Cm.Hormoligosis , the stimulatory effect of harmful agentswhe n applied at sub-harmful doses,coul d account at least inpar t for this improved fecundity (158).

36 Theus eo fplan tproduct st oprotec tstore dseed s

4.1.3EFFECT SO N EGGSAN D LARVAE Thejuvenil e stages ofth e storage insects are generally less affected byvolatil e oils than the adultbeetles ,bu t they areusuall y not completely tolerant to thetreatments .Egg so f Ccwer e susceptible to vapours ofAcorus calamus. Th eyounge r embryonic stages were more susceptible thanth e later stages (278)bu tth e oil didno t affect larvae and pupae f264). For Cm,th e fertility ofegg swa sheavil y affected bytreatmen t with Cymbopogon schoenanthusan dLavandula spp.(106) . Ocimum basilicuman dPelargonium spp . showed highest effect onreductio n of egghatchin g and on adult emergence for Cc (263).Piper guineense prevente d adult emergence of Cmcompletel y (216).Pee l oilso f eight citrus species all showed adecreas e innumber s ofprogen y of Cmfro m treated beans (323).Number s ofoffsprin g ofSg, So an d Ccemergin g from seedstreate d with Acorus calamusoi lvapour s were considerably lower than inth e controls.Th e response was correlated to increase ofexposur e timerathe r thant o increase in dose(279) .

4.2 COMPARISON OFBEETL E SPECIES Beetle species differ in susceptibility to volatile oils.Th e vapours ofAcorus calamus weremor e effective against Cctha n against Sg andSo. Adults ofbot h Tcoan dRd wer e completely tolerant (87;264 ;278) .Eucalyptus citriodora an d Ocimumbasilicum wer e not toxict oSo bu t werepoten t against Cm(96) . Coriandrum sativum wastopicall y non­ toxict o Cm,Ls and Tco, butth e oilwa srepellen t andmoderatel y toxict oSo and repellent to Tco(320) .Citru spee l oils showed more effect against adults of Cmtha n against adults ofD m andSz (79).Topica l application of eight lyophilised citruspee l oils was highly toxic to Cman d moderately toxic toSo adults(324) .

4.3 COMPARISON OFPLAN T SPECIES Not all volatile plant oils are equally effective asprotectant s of stored products.I nth e reviewed articles,th e most effective plants found incomparison s areofte n ofth e family ofth e Lamiaceae. Coleusamboinicus wasmor e effective asa contact toxicant against Ao and Cctha nAgeratum conyzoides, Blumeabalsamifera, Chrysanthemum indicum, and Vitex negundo(199) .Al l oils ofLamiacea e tested by Djibo etal. (72 )wer e toxict o adults of Cm;Ocimum basilicum wa s three times moretoxi c thanHyptis spicigera an d fourteen times moretoxi c thanHyptis suaveolens. Ocimum basilicumwa smor e toxic than Tagetes minutaan dPiper nigrum to eggs and adults of Cm(141) . For Cymbopogon, arankin g inth e degree ofeffectivenes s isno t evident. Bhaduri etal. (36) showed that Cymbopogon citratusan d Cymbopogon narduswer emor e effective than seven non-volatile oilsagains t ovipositiono f Cm.Adebay o and Gbolade (2) found that Cymbopogon citratus,Eugenia uniflora, Lantanacamara an dLippia adoensis completely inhibited oviposition and adult emergence of Cm,wherea sEupatorium odoratumwa sles seffective . Cymbopogon schoenanthusan dLavandula spp.wer ever y toxic to Cm,causin gparalysi s and death at low dosages,whereas , onth eothe r hand Cymbopogon citratus, Cymbopogon nardus,Cedrus spp. ,Diplolophium africanum, Eucalyptuscitriodora an dLippia multiflora were less toxic (106).Rich a etal. (263 ) found that Ocimumbasilicum an dPelargonium spp.wer e more effective in causing adult mortality of Cctha n Cymbopogon citratus,Cymbopogon nardusan d Galega spp. whereas Citrus limonan dEucalyptus spp .wer eno t effective atall . Forth e family ofth e Rutaceae, some contradicting results havebee n found. Citrus aurantifolia,Citrus limon, Citrus paradisi and Citrus reticulatacause d complete adult mortality for Cm,wherea s Citrus sinensisan d CitrusX tangelower e not toxic(324) .

37 4Volatil eoil s

Citrusparadisi completely inhibited egghatc h ofCm whil e Citrussinensis was less effective (89).However , Don-Pedro (73)foun d that Citrussinensis wa smor e effective against Cman dDm adults than Citrusparadisi and Richa etal. foun d that Citrus limon hadn oeffec t onmortalit y of Cc (263). Acorus calamus isofte n tested, but never compared to other oils.Acorus calamusoil s from three origins differed inefficacy , but athig h doses they were all completely toxic to adults ofCph (246) .

4.4DURATIO N OFTH E EFFECTS Contradictory results havebee n found for the duration of theeffec t ofvolatil e oils.Th e oilsar e often found effective for only ashor ttime ,wit h no residual action. Citruspee l oil treatment caused 100%mortalit y of Cm andDm adults at onehou r after application, butth e oil lost all activity within 24hour s and didno t show anyresidua l activity on eggs orlarvae .Th eactivit y of lime-peel oil was dependent on thetim e interval between application ofth eoi lan dth e start ofth ebioassa y (77).Dennettia tripetala an dPiper guineenseachieve d complete adult mortality within onewee k but these and other (Xylopia aethiopicaan dMonodora myristicd) volatile oilswer eno t effective as grain protectants because they did allowth e development of anF l generation (215). However, for somevolatil e oils alonge r lasting effect hasbee n found. Piperguineense (216)Cymbopogon citratusan d Cymbopogon narduswer eeffectiv e against Cmfo r up to 90 days (36). Citrusparadisi and Citrus sinensisretaine d their effect upt o 150day s after treatment (89).Acorus calamus showe d alon g lasting residual effect and could ensureth e storage roomt ob e free from pulsebeetle s for many months (351). Emergence of Cmfro m seeds treated with peel oils ofeigh t citrus species was lowunti l 312 days after treatment (323). Generally, theduratio n of exposure ofth ebeetle s to the oil ismor e important than the applied dosage ofth e oil asa factor affecting theefficienc y (87).Th e response after treatment withAcorus calamuswa s correlated to an increase ofth eexposur e time rather than toa n increase indos e (279).Thi s was indicated by the increase ofbeetl e mortality after an increasing period ofexposur e at acertai n dosage (278).Th e LC5ofo r Callistemon citrinusan dEupatorium capillifoliumfo r adults of Cmdecrease d with exposure time(25) .

4.5 EFFECTS ONTH E STORED PRODUCT The effects ofvolatil e oilso n stored seeds areminima l because the oilsusuall y dono t need tomak e contact with the seedst ob e effective. None ofth e investigated preparations ofplan t origin had anegativ e influence onwhea t seed germination (138). Formulations ofAcorus calamuswer enon-hazardou s anddi dno t impair seed germination (351).Edibl e volatile oils couldb ever yusefu l on farm level in developing countries aspotentia l control agents against storage beetles. They could play an important role instored-grai n protection, reducingth enee d for, andris k associated with, theus e of insecticides (293).

38 Theus eo fplan tproduct st oprotec tstore dseed s

4.6 CONCLUSION Volatile oils are effective against storage beetlesbu tthei r period of action isusuall y not long because they quickly evaporate unless airtight structures areuse d for storage. Volatile oils mainly affect adult beetles.Th e effect canb e eitherrepellen t ortoxic ,bu t duet o laboratory set-ups thetoxi c effect isofte n overestimated. Through the adults, oviposition isaffecte d and oilvapour s can affect larvae andeggs . Application iseasy ,bu t should berepeate d for-long termprotection . The insecticide doesno tusuall y need totouc hth e stored product, soth eproduc t ismostl y unaffected by the treatment. Toobtai n large enough quantities ofvolatil e oil, large quantities ofplants , distillation equipment and alo t ofwor k are needed.

39 4Volatil eoil s

40 Theus eo fplan tproduct st oprotec tstore dseed s

5NON-VOLATIL E OILS Non-volatile oil,use d asa coatin g for seeds,ca n effectively protect these seeds against insect pests during storage.Th efilm o f oilprevent s the attachment ofth e eggt oth e seed coat andplug s the respiratory systems ofegg s andadul tbeetles . Oilsca nb e extracted mechanically orb y hand from seeds cheaper than the stored product or from seedso f non-crop plants. To obtain enough oil for treatment, large quantities ofplan t material are needed andth e extraction and application ofnon-volatil e oils areno t easily done. However, most ofth e oils are very effective and retain their effectiveness over alon g period. Allplant s discussed inthi s chapteran dliste d intabl e 5ar euse d for theirnon ­ volatile oil unless stated otherwise.

Table5: Plants of which the non-volatile oil is used against storage insects, the quantity ofoil usedand its effect. Plant (sub-) Plant species Quantity - Beetle species A Affected stage # Refe­ family A O E H L M P rence

Arecaceae Cocos nucifera 1-20 Ao, Cc *! *! 199 10 Ao, Cm !! ü ** 271 1-4 Ca *! *! *! 157 5-10 Cc il il n M il 11 3 Cc ** *! *i 32 0.5-3 Cc *! 33 10 Cc ü 127 10 Cc *! 128 0.5% Cc *i 190 50g/kg Cc *! 193 0.5-5 Cc *! 201 1-4 Cc *i 294 2-4 Cc *! *! 295 5-7.5 Cc *i 298 1-5 Cc *! *! *i 299 1-3 Cc *! *! 306 l-8,3g/kg Cc, Cm *! 111 150 ng/bean Cc, Cm *! 352 1-10 g/kg Cc, So *! *! *! *! 292 3% Cc, Sz *! 200 2-8 g/kg Cm *! 36 3.5-14 Cm ** *! *! 74 1.75-14 Cm *! *! 75 Dipped Cm ** ** *! 76 5 Cm *! *! *i *! 181 2.5-10 Cm ** ** *! 203 2.5-10 Cm ** ** 204

41 5Non-volatil eoil s

Plant family Plant species Quantity Beetle A O E H L M P Ref. Arecaceae Cocos nucifera 1.5g/kg Cm *! *! 256 Continued 0.4-2.5 g/kg Cm *! M M 290 1-8 Cm *! *i 305 5-10 Cm *! 341 4-8 g/kg Cr *i 250 1-5 Zs *! 283 Arecaceae Elaeis guineensis 5-10 Cc M H *! *! M 11 10 Cc *! 128 5-10 Cc *! ** 136 5-10 Cc *i *! 144 5-10 Cc ** ** 145 10 Cc *! 146 2-4 Cc ** *! 295 2-8 Cc ** H H 342 150ng/bean Cc, Cm *! 352 1-10 g/kg Cc, Cm, So *! ** *! *! 292 1-8 Cm *i 111 2.5-10 Cm ** ** *i 203 2.5-10 Cm *i ** 204 0.25-8 Cm *! *! ** 235 5 Cm *! 268 0.4-2.5 g/kg Cm *! M n 290 1.5-3 g/kg Cm *i 293 1-8 Cm *! *! 305 2.5-30 Cm *i *! *! 334 5 Cm *! 341 6 Cm, Rd, So, Tea, Tg *! 8 1 Zs *i *! *i 118 1-5 Zs M M M 283

Asteraceae Carthamus tinctorius 5-40 ml/1 Cc *! *! 5 5-10 Cc *! *! 136 5-10 Cc *! *! 144 5-10 Cc ** •* 145 10 Cc *! 146 2.5-10 Cc ** *! *! 272 1-3 Cc *! *! 306 1-10 g/kg Cc.So *! ** *! *i 292 5 Cm *! *! *! *! 181

42 Theus eo fplan tproduct st oprotec tstore dseed s

Plant family Plant species Quantity Beetle A O E H L M P Ref. Asteraceae Carthamus tinctorius 2.5-10 Cm ** ** *! 203 continued 2.5-10 Cm ** ** 204 0.4-2.5 g/kg Cm *! ü M 290 Asteraceae Guizotia abyssinica 2.5-10 Cm ** *! *! 203 2.5-10 Cm *! ** 204 Asteraceae Helianthus annuus 5 Ao *! 338 1-4 Ca ** ** ** 157 5-40 mVl Cc *! *! 5 5-10 Cc *! *! 136 5-10 Cc *! *! 144 5-10 Cc *• ** 145 10 Cc *i 146 0.5-5 Cc ** 201 2.5-10 Cc ** *! M 272 2-4 Cc *! *! 295 1-3 Cc *! *! 306 5-10 Cc, Cm, Cr *! *! 251 1-10 g/kg Cc, So *! *i *i *! 292 3g/k g Cm *! 111 2-30 Cm *! 255 0.4-2.5g/k g Cm ** *! *! 290

Bombacaceae Ceiba pentandra 1-10 g/kg Cc, So ** ** *i *! 292 0.4-2.5g/k g Cm ** ü H 290

Brassicaceae Brassica juncea 1-4 Ca ** ** ** 157 5-10 Cc *! ** *i *! *! 11 5-10 Cc *! *! 144 1-4 Cc *! 294 2-4 Cc ** *! 295 1-5 Cc ** *! *! 299 1-3 Cc *! il 306 2-8 Cc ** ü M 342 150 ng/bean Cc, Cm M 352 Brassicaceae Brassica spp. 1-4 Ca ** ** ** 157 5-10 Cc *! ** u *! *! 11 3 Cc *! *i *! 32 0.5-3 Cc *! 33 5-10 Cc *! *! 136

43 5 Non-volatile oils

Plant family Plant species Quantity Beetle A O E H L M P Ref. Brassicaceae Brassica spp. 5-10 Cc ** ** 145 continued 10 Cc *! 146 5-10 Cc H n 148 0.5% Cc *! 190 50g/kg Cc M 193 0.5-5 Cc *i 201 0.25-1% Cc *! *! *! 231 2.5-10 Cc *! 260 2.5-10 Cc *! *! n 272 1-4 Cc *! 294 2-4 Cc ** *! 295 5-7.5 Cc *! 298 1-5 Cc *! *! *! 299 15 Cm n 6 2-8 g/kg Cm *! 36 2.5, 3g/k g Cm *! 111 2.5-10 Cm ik* *! *! 203 2.5-10 Cm *! ** 204 2-30 Cm u 255 Brassicaceae Eruca vesicaria 2-4 Cc *! *! 295 1-3 Cc *! H 306 Brassicaceae Raphanus sativus 1-3 Cc ** *i 306

Clusiaceae Calophyllum inophyllum 1-3 Cm ** *! H n 129 2.5-10 Cm ** *t n 203 2.5-10 Cm *! ** 204 Dipterocarpacea« Shorea robusta 3-5 g/kg Cm *! 111 3-5 Cm ** 229

Euphorbiaceae Jatropha curcas 1-3 Cm ** *! H H 129 Euphorbiaceae Jatropha indica 10 Cm, Sz n 22 Euphorbiaceae Ricinus communis 10 Ao, Cm M M ** 271 5-10 Cc *! M 136 5-10 Cc n *i 144 5-10 Cc ** *! 145 10 Cc *! 146 0.5-5 Cc *i 201 2.5-10 Cc *! 260 2.5-10 Cc *! *! M 272

44 Theus eo fplan tproduct st oprotec tstore dseed s

Plant family Plant species Quantity Beetle A O E H L M P Ref. Euphorbiaceae Ricinus communis 1-3 Cc *! u 306 continued 5-15 Cc M n *! n 355 2-8g/kg Cm »! 36 1-8 Cm *! 111 2.5-10 Cm »* *! n 203 2.5-10 Cm *! ** 204 1-8 Cm *! *i 305 5-10 Cm, Cph *! n 222 4-8g/kg Cr *! 250

Leguminosae- Senna occidentalis 10 Cm *! *! *i 165 Caesalpinioideae 10 Cm *! *! 180

Leguminosae- Arachis hypogaea 10 Ao, Cm n \\ ** 271 Papilionoideae 5-10 Ba, Cm *! *! 171 1-4 Ca *! *! *i 157 3 Cc ** *! *! 32 0.5-3 Cc *! 33 5-10 Cc *! n 136 5-10 Cc *i *! 144 5-10 Cc ** ** 145 10 Cc *! 146 0.5% Cc *i 190 50g/kg Cc u 193 0.5-5 Cc *t 201 2.5-10 Cc ** 260 1-4 Cc *i 294 2-4 Cc ** *i 295 5-7.5 Cc *t 298 1-5 Cc *! *i *! 299 1-3 Cc *i *! 306 2-8 Cc ** n n 342 l-10,3g/kg Cc, Cm *! n 111 150 u^ean Cc, Cm *! 352 5-10 Cc, Cm, Cr ** *! 251 1-10 g/kg Cc, So »* *! *! *! 292 5-10 Cm H 17 5 Cm *i 20 2-8 g/kg Cm *! 36 Flour pellets Cm *i *! u *i 42

45 5Non-volatil eoil s

Plant family Plant species Quantity Beetle A O E H L M P Ref. Leguminosae- Arachis hypogaea 5 Cm *i 52 Papilionoideae continued 10 25 g/kg Cm *! *! *! *! 60 10 Cm *! *! *! 61 3.5-14 Cm »* *! *! 74 1.75-14 Cm *! *! 75 Dipped Cm ** ** *! 76 7 Cm ** *! *! 77 1-5 Cm *i *! *i *! 181 2.5-10 Cm ** ** *! 203 2.5-10 Cm ** ** 204 5-6 Cm H 217 0.25-8 Cm *i *! ** 235 5 Cm *i 239 2-30 Cm *! 255 0.4-0.8g/k g Cm ** *! *! 290 1-8 Cm n *! 305 2.5-30 Cm *! *i n 334 1-5 Cm *! 341 5 ml/1 Cm, Tco *! 354 10 Cm, Sz M 22 4-8 g/kg Cr *! 250 Leguminosae- Glycine max 5 Ao *! 30 Papilionoideae 10 Ao, Cm !! n ** 271 1-4 Ca ** ** ** 157 3 Cc ** *i *! 32 0.5-3 Cc *! 33 5-15 Cc n 57 0.25-1% Cc *i *i *i 231 1-4 Cc *! *! ** 265 2-4 Cc ** *! 295 1-5 Cc *! *! *! 299 1-5 Cc *! 303 1-3 Cc ** n 306 2-8 Cc *• n M 342 150ng/bean Cc, Cm «* 352 1-10 g/kg Cc, So *! ** *! *! 292 2.5-10 Cm ** *! *! 203 2.5-10 Cm ** ** 204

46 Theus eo fplan tproduct st oprotec tstore dseed s

Plant family Plant species Quantity Beetle A 0 £ H L M P Ref. Leguminosae- Glycine max 2-30 Cm *! 255 Papilionoideae continued 0.4-2.5 g/kg Cm ** M M 290 5-10 Cm, Cph *! 1! 222 4-8g/k g Cr *! 250 1-5 Zs *! *! *! 283 Leguminosae- Pongamia pinnata 5-10 Cc *! H 136 Papilionoideae 5-10 Cc H *i 144 5-10 Cc ** *! 145 10 Cc *! 146 10-200 Cc *! *! 206 2.5-10 Cc *! 260 2.5-10 Cc *! *! H 272 2-8 g/kg Cm H 36 2.5-10 Cm ** »* M 203 2.5-10 Cm H ** 204

Linaceae Linum usitatissimum 1-4 Ca ** ** ** 157 5-40 ml/1 Cc *! *! 5 10 Cc ** 111 0.5-2 ml Cc ** 184 1-3 Cc ** *! 306 10 Cc, Cm *! *! *! *! *! *! 245

Malvaceae Gossypium hirsutum 1-4 Ca *! *! *! 157 2.5-10 Cc *! *! H 272 1-4 Cc *! 294 2-8 Cc *! M !! 342 5-15 Cc M *! ** *! 355 150 ug/bean Cc, Cm *! 352 1-10 g/kg Cc,So *! ** *! *! 292 10 Cm *! *! *! *! 60 10 Cm ** *! 61 3-5 g/kg Cm *! 111 2.5-10 Cm ** »* *! 203 2.5-10 Cm ** ** 204 3-5 Cm *i 229 2-30 Cm *! 255 0.4-2.5 g/kg Cm ** M M 290

47 5 Non-volatile oils

Plant family Plant species Quantity Beetle A O E H L M P Ref. Malvaceae Gossypium hirsutum 1 Zs *! *! *! 118 continued 1-5 Zs Ü *! n 283

Meliaceae Azadirachta indica 5-100 g/1 Ao, Cc *! *! 199 2.5-7.5 Ao, Zs n 47 1-3 Ca, Cc, Cm ** *! 282 1-10 Ca, Cc, Cm, Rd, Sz, *! *! *! 275 Tea, Tco 5-40 ml/1 Cc *! n 5 5-10 Cc it il n 1! n 11 10 Cc u 64 10 Cc *! 128 5-10 Cc n {! 136 5-10 Cc n *! 144 5-10 Cc ** *t 145 10 Cc *! 146 0.5-2 ml Cc *i 184 10-50 g/kg Cc *! 228 1-3% Cc *! ** 244 2.5-10 Cc ** 260 2.5-10 Cc *! *! M 272 10 Cc, Cm H *! *! n n *! 245 3% Cc.Sz *! 200 2-8 g/kg Cm *! 36 25 g/kg Cm *i *! *! 60 1-10 Cm !! n n 63 1-3% Cm *! 111 2-3 Cm *i *! *i 124 1-3 Cm ** *! n 129 2.5-20 g/kg Cm n 161 2.5-10 Cm ** *! 203 2.5-10 Cm n ** 204 0.25-8 Cm n *! *! 235 2% Cm ** 255 1.5 g/kg Cm *! *! 256 10 Cm.Sz u 22 5 ml/1 Cm, Tco *! *! *i 354 4-8 g/kg Cr *! 250 Meliaceae Melia azedarach 2-8 Cc *! *i 356

48 The use of plant products to protect stored seeds

Plant family Plant species Quantity Beetle A O E H L M P Ref.

Oleaceae Olea europaea 10 Ao, Cm H H ** 271 5-15 Cc it *! *! n 355 150 (xg/bean Cc, Cm *! 352 1-10 g/kg Cc,So *! *! *! *i 292 15 Cm *! 6 0.4-2.5g/k g Cm ** H n 290 2.5-30 Cm ** 334 10 Cm *! 341

Pedaliaceae Sesamum indicum 10 Ao, Cm M H ** 271 1-4 Ca *! *! ** 157 5-40 ml/1 Cc *! *i 5 5-10 Cc it il il *! M 11 10, 3 g/kg Cc *i 111 10 Cc *! 128 5-10 Cc *! *! 136 5-10 Cc *! *! 144 5-10 Cc ** ** 145 10 Cc *! 146 0.5-2 ml Cc ** 184 0.5-5 Cc *! 201 0.25-1% Cc *t *! *! 231 1-4 Cc *! 294 2-4 Cc *! *! 295 1-5 Cc *! *! *i 299 1-3 Cc ** *! 306 5-15 Cc n *i *! n 355 150 ng/bean Cc, Cm *! 352 5-10 Cc, Cm, Cr ** *i 251 2.5-10 Cm ** *! *! 203 2.5-10 Cm ** ** 204 15-20 Cm *i 341

Poaceae Oryza sativa 1-5 Cc n 45 1-3 Cc *! *! 306 1-8 Cc *i il 353 150 ng/bean Cc, Cm *! 352 1-10 g/kg Cc, Cm, So ** *! M Ü 292 3-5 g/kg Cm *t 111

49 5Non-volatil eoil s

Plant family Plant species Quantity Beetle A O E H L M P Ref. Poaceae Oryza sativa 3-5 mg/kg Cm *! 229 continued 0.4-0.8 g/kg Cm ** M ij 290 5 Cm *! 341 1.5-10g/k g Cm, Sz *! 293 Poaceae Zea mays 5 Ao *! 338 5 Ao, Zs *! *! 309 5-10 Cc *! 57 5-10 Cc *! *! 136 7.5-12.5 Cc *! *! 144 5-10 Cc ** ** 145 10 Cc *! 146 5-15 Cc Ü H M i; 355 5-10 Cc, Cm, Cr ** *! 251 1-10 g/kg Cc,So *! *! *! *! 292 0.4-2.5 g/kg Cm *» !! !! 290 1-5 Zs *! *! *! 283

Ü Sapotaceae Madhuca longifolia 5-10 Cc •*! !! ** !! 11 2.5-10 Cc *! *! î! 272 2-8 g/kg Cm *! 36 1-3 Cm ** ** *! H 129 5 Cm *! *! 247 1.5 g/kg Cm *! *! 256 4-8 g/kg Cr *! 250 Sapotaceae Vitellaria paradoxa 10 Cm *! *! *! *! 60 10 Cm ** *! 61 0.25-8 Cm *! *! ** 235

Dalda 5-10 Cc *i *! *! *! il 11 ~:Quantitie si nm loil/k gstore dproduct ,unles sstate dotherwis e A:Se etabl e1 #:A =adul tlongevit y& fecundity , O =oviposition ,E = surviva lo fegg s& embryoso nth esee d surface, H= hatching ,L =surviva lo flarva ean dpupa einsid eth eseed ,M = emergence ,P = populatio n numbersi effect onth estore dproduct ;* *= Measured ,bu tn ostatisticall ysignifican t resultswer efoun do r presented, *! =Significan t decrease, !! =Tota linhibition .

5.1 DEVELOPMENTAL STAGE AFFECTED

5.1.1 EFFECTS ON ADULTS Non-volatile oilsca nhav enegativ e effects on adult beetles,eithe rthroug h contact toxicity orthroug h deterrence.Al l oils tested by Salas andHernande z (271) caused complete adult mortality forAo and Cm.Arachis hypogaeacause dhig h adult mortality for Cm(217) .Arachis hypogaeaan dElaeis guineensis decreased adult longevity ofCm

50 Theus eo fplan tproduct st oprotec tstore dseed s

(334).Elaeis guineensis increased adult mortality andreduce d reproduction ofZs. The insecticidal effectiveness was determined by thetriglycerid e component ofth e oil(118) . Cocos nucifera(127) ,Madhuca longifolia, Sesamumindicum, Azadirachtaindica, an d Elaeisguineensis inflicted complete adult mortality for Ce(11). Glycine max showed high efficacy against eggs andadul t stages of Cc(265) .Al l oilsteste db y Zewar (355) caused mortality of Ccadults .

5.1.2 EFFECTS ON OVIPOSITION Oviposition can be influenced by treatment ofth e stored product withnon-volatil eoil . Thedecreas e inth e number ofegg s couldb e due to other causestha njus t theeffec t on the longevity ofth e adult beetle.Ricinus communis, Arachis hypogaea, Pongamia pinnata andAzadirachta indica affected oviposition and induced egg mortality for Cc (136). Seedtreatmen t withAzadirachta indica (64), Cocosnucifera (199) ,Madhuca longifolia,Sesamum indicum an dElaeis guineensis prevente d adults ofCc fro m laying eggs (11).Zea mays, Arachis hypogaea, Helianthusannuus an dSesamum indicum reduced oviposition of Cm,Cc an d Cr(251) .Al l oilsteste db y Sangappa (272)an db y Singal and Singh (299) reduced oviposition of Ccsignificantl y and the oils tested by Singh etal. (306 ) didth e same for Cm.O nbambar a groundnut,Azadirachta indica, Elaeisguineensis, Arachis hypogaeaan d Vitellariaparadoxa had adecreasin g effect on oviposition of Cm(235) .Azadirachta indica(161) , Cocos nuciferaan dMadhuca longifoliadeterre d oviposition of Cm(256) . Jatrophacurcas, Azadirachtaindica, Calophyllum inophyllum andMadhuca longifolia al lpartl yprevente d oviposition (129). Oviposition of Cmwa s completely inhibited after seed treatment withPongamia pinnata (36).

5.1.3 EFFECTS ON EGGSAN D LARVAE Most ofth e efficacy ofnon-volatil eoil s canb eattribute d toth e effect on eggs andthei r attachment to the seed coat. The effect ofth e oils isa t leastpartl y based on mechanical action. Due toth e oil layer around treated seeds,egg s cannot be attached effectively to the seed surface. Hatching offirst insta r larvae isprevente d becausepenetratio n ismor e difficult ifeg g attachment isles s secure (75).Fo r Cm,th e funnel through which the larvae should hatch into the seed, is clogged by the oil soth e larva cannot hatch and dies inth e egg stage(55) .If th eeg gha d already been laidbefor e treatment ofth e seed, the oilwil l plug itsmicropyle ,whic h causesth e embryo insideth e egg to die from oxygen starvation (202).Th e oil could alsopenetrat e the chorion ofth e egg via the micropyle andcaus e death ofth edevelopin g embryothroug h asphyxiation (161).Arachis hypogaeaha d anovicida l and suffocating effect onbruchids ,wit h eggs and young larvaebein g more sensitive than older larvae and nymphs (171).Egg s of oneo rtw o days oldwer e killed immediately, whereas inthre e to five day old eggs deatho fth e developing larvae occurred inminute s(17) . In addition to the mechanical effect, acombinatio n ofmechanica l andtoxi c effects could occur. The ovicidal effect ofoil s ofArachis hypogaeaan d Cocos nuciferacoul d be caused by a direct toxic action after penetration of theoi l intoth e egg. However, the effect could also be caused by thereductio n ofrespirator y activity ofth e egg andb y accumulation oftoxi c metabolites inside ita sa resul t ofth e oil barrier effect (75). Effects being solely toxichav eals obee n found. Oryzasativa, Shorea robusta an d Gossypium hirsutumentere d the eggthroug h the micropyles, and thetoxi c components ofth e oils destroyed theyol k (229). However, thewa y in whichth e effect isaccomplishe d has not always been elucidated. The insecticidal activity ofnon-volatile , edible oilsan do fth e active fatty acids

51 5Non-volatil eoil s

extractedfrom them , depended mainly on their ovicidal effect (74; 76).Fo rCm, Carthamus tinctorius,Cocos nucifera an dArachis hypogaeacause d high mortality of eggs andlarva e onth e seed surface, butha d no effect onindividual s that successfully entered the seed. The lack ofoil-specifi c activity indicated the effect tob e physical rather than chemical (181).Azadirachta indica an d its extract effectively reduced egg hatching of Cm(203) .Fo r Cc, Ricinuscommunis, Arachis hypogaea, Pongamiapinnata andAzadirachta indicacompletel y prevented hatching ofth e eggs (136). Jatropha curcas, Azadirachta indica,Calophyllum inophyllum andMadhuca longifoliaprevente d egghatchin g and emergence (129).

5.2 COMPARISONS OFBEETL E SPECIES The effects of different oils can vary whenthe y are tested on different beetle species. Oryza sativa, Glycine max, Gossypium hirsutuman dElaeis guineensis, were potential control agents against Cm,bu t they were less effective against Sz and So(293) . Zea mays, Helianthusannuus, Arachis hypogaea, andSesamum indicum significantly reduced adult longevity of Cman d Cc,wherea s only the latter two oils affected Cr (251).So was best controlled with refined Elaeisguineensis, wherea s Tea and Tgwer e most sensitive to fresh Elaeisguineensis and Cmwa s susceptible tobot h (8). For effective control ofSo and Sz,dosage s of oilneede d are higher than for the control of Cm (292).

5.3 COMPARISON OFPLAN T SPECIES If different oilsar e compared, a few general conclusions can be drawn. - Crude oils are more effective than refined ones from the sameplant . Fresh oil ofElaeis guineensis wasmor e effective than bleached and refined oil (8).Agains t Cc, purified Gossypium hirsutumwa s not effective (355).Crud e oils of Gossypium hirsutum, Glycinemax an d Cocosnucifera wer e more effective than refined ones (283).Crud e oilsteste d by Shaaya andKostjukovsk y (292) mostly resulted in fewer eggs laid and fewer emerged adults of Cctha n after treatment with refined or distilled products made ofthes eoils . -Non-edibl e oils seem tob e generally moreeffectiv e than edible oils (203).Ricinus communiswa s more effective than Glycine maxagains t Cman d Cph(222) .Amon g the non-edible oils,Azadirachta indicaoi li s often one ofth e most effective ones. Azadirachta indicawa s most toxic to adults of Ccwhe n compared with Cocos nucifera (199) orLinum usitatissimum (245) .Azadirachta indicagav e complete protection against Cc(64 ) and ofal l tested oils,Azadirachta indica wa s the only onetha t completely prevented oviposition of Cmo n cowpea (235). Azadirachta indicaan d its extract were moreeffectiv e thanRicinus communis, Pongamiapinnata and Calophyllum inophyllum, which themselves weremor e effective than the edible oilso f Sesamumindicum, Brassica spp., Guizotia abyssinica,Gossypium hirsutum,Elaeis guineensis,Arachis hypogaea,Cocos nucifera,Carthamus tinctoriusan d Glycine max (203). Different mechanisms seem tob e involved.Helianthus annuus wa s effective against oviposition of Cc,wherea s Glycine max,Brassica juncea, Arachis hypogaeaan dOryza sativaprotecte d against adult emergence andEruca vesicaria, Cocos nucifera, Carthamus tinctoriusan dRicinus communis wer eeffectiv e against both(306) . - When edible oils are compared, crude cotton (Gossypiumhirsutum), an d Brassicaceae areusuall y among the most effective ones. Gossypium hirsutumwa s more effective

52 Theus eo fplan tproduct st oprotec t storedseed s against Cmtha n Oryzasativa an d Shorearobusta (229) .Brassica spp.wa s more effective than Oleaeuropaea (6).Arachis hypogaea,Cocos nucifera, Brassica spp. , Sesamumindicum, Glycine maxan dBrassica juncea were all effective against Cc. Brassica spp.wa s most effective (299). Arachis hypogaeaan dBrassica spp.wer e more persistent than Cocosnucifera (193). For a few oils,contradictor y resultshav e been reported. Both fresh andrefine d oilso f Elaeisguineensis wer e effective against Cman dRd (8).Elaeis guineensis was the most effective crude oil(283) ,mor e effective than Zeamays and Arachis hypogaea(151) . However,Elaeis guineensis wa s found to beth e least effective of 10oil steste d by Khaire etal. (144) , inthi s article itwa s found to be less effective than, among others, Zeamays. Emergenc e wasprevented , indecreasin g order of effectiveness by (non- edible)Azadirachta indica, Pongamiapinnata, Ricinus communis, an d seven edible oils (144). However, in other experiments Pongamiapinnata was more effective than Azadirachta indica (272),Ricinus communis an d four edible oils (36).Accordin g to Singh (305),Arachis hypogaeacompletel y prevented emergence of Cm whereasRicinus communis,Cocos nucifera an dElaeis guineensis onl y caused partial reduction but Pierrard (239) found thatArachis hypogaeafaile d to completely control Cm.

5.4 DURATION OF THE EFFECTS The time scale ofth e effect aimed for isno t the same in every article. Some authors aim for aprotectio n mechanism that works immediately, whereas others intend to keepth e level of damage inth e stored product below acertai n threshold for as long aperio d as possible.Fo r the immediate knockdown effect on adultbeetles ,non-volatil e oils are usually less suitable. Non-volatile oils seem to be most effective for intermediate periods of protection, weeks to a few months. In the Sahel,oil s diminished infestation only temporarily, whereasi n other, more humid regions the method appeared tob e effective (20).Th e protection offered by oils ofAzadirachta indicaan dPongamia pinnata lasted almost twice aslon g astha t ofedibl e oils (204).Pongamia pinnata completely inhibited oviposition, but its efficacy sharply deteriorated at later stages (>30 days) (36).Ricinus communis, Pongamiapinnata andAzadirachta indica completel y protected stored seeds against Cc for 33 days and partly upunti l 100days .Arachis hypogaea prevente d egg hatching beyond 33 days (136).Linum usitatissimum andAzadirachta indicawer e effective for three and four months respectively (245).Th e toxic effect of Cocos nuciferaagains tCc lasted for at least 10week s (127).Th e effect ofArachis hypogaeaagains t Cmpersiste d for 3month s (217). Oryzasativa gav e complete protection against Ccfo r upt o 135day s (45).Azadirachta indicaeffectivel y protected gram seeds against bruchid damage for at least 135day s (228).If th etreatmen t was repeated after 12day s and after 4month s of storage, Vitellariaparadoxa was ablet o effectively protect seeds stored injar s or sacks (153). Other authors waited longer and found the effects to last even longer.Ricinus communis protected stored cowpea for up to 150days . Glycine maxreduce d the population build up for up to 90day s (222).Wit h Gossypium hirsutum,infestatio n started at the same time aswit h Oryza sativa andShorea robusta, but itremaine d almost negligible up until 5 months (229). Cocosnucifera protecte d beans for 4 months against insect infestation andArachis hypogaeaan dBrassica spp .di d so for 6month s (190; 193). Azadirachta indicagav e complete protection against Ccfo r at least 5month s (64). Azadirachta indica(63) ,Jatropha indicaan dArachis hypogaeacompletel y controlled Cman dSz for 6 months (22). Seeds treated withArachis hypogaeacoul db e stored for 6t o 12month s

53 5 Non-volatileoil s

without beetle attack (171). Glycine maxcause d damage to remain below the economic threshold for 8month s (30).Pongamia pinnata protected storedre d gram for 319day s and remained active afterwards (272).

5.5 EFFECTS ONTH E STORED PRODUCT Disadvantages exist ifoil s are used for long-term storage of seeds.Zea mayscause d a change of colour inth e seeds (309).Product s of Vitellariaparadoxa became rancid and could therefore beuse d for sowing seeds only (66). Azadirachta indicaan d Ricinuscommunis ha d anegativ e influence onth etast e ofth ebeans .Azadirachta indicaan d Guizotia abyssinicaha d anegativ e effect on seed germination after 6 months of storage (204).Seed streate d with Cocos nuciferao rwit hArachis hypogaea failed to germinate (334).Wate r absorption by stored seedswa s affected after oil treatment (355) andth e cooking time of oil-treated seeds wasprolonge d (231). Another disadvantage isth e difficulty of applyingnon-volatil e oilsespeciall y for greaterquantitie so fbeans .I fth efilm aroun d thebea n doesno t completely coverth e surface, the efficacy ofth e oiltreatmen t will bemuc h lower since the mechanical effect ofoil s isusuall y theke y tothei r efficacy. Acomplet e film over the seed surface isvita l for the success ofnon-volatil e oils asprotectiv e agents of stored products (75). Aunifor m spread ofth e oil onal l grain would maximise the effect (74). Advantageous effects werereporte d aswell .N o effect on seed germination was found after treatment with either Glycine maxo rZea mays fo r 8month s (57) orwit h Helianthusannuus fo r sixmonth s (338), and thepalatabilit y and appearance ofth e beans improved duet othei r shiny coating.

5.6 CONCLUSION Non-volatile oilshav emainl yphysica l effects. They are most effective ifthe y completely cover the surface of thestore d seeds and therefore care should betake n to applythe mproperly . The mechanical action seems tob e most effective againsteggs , whereas adultsar e more affected byth e chemical action. Larvae andpupae , as they are insideth ebean , are generally less sensitive to the effects of oils.Th eeffect s ofoil s onth e stored product are numerous,rangin g from a slight change ofcolou r to asever e change oftast e and inhibition of seed germination. To obtain large enough quantities of oil,grea t quantities ofplan t material are needed and lots ofwork . From onekil oo f dryAzadirachta indicakernel s 75-100 ml of oil can beobtaine d byhan d (21).Mos t ofth e edible oils areusuall y for sale.

54 Theus eo fplan tproduct st oprotec t storedseed s

6 EXTRACTS Extracting plant material with an appropriate solvent generally results in concentration ofactiv e ingredients. Such extracts aretherefor e often moreeffectiv e against storage beetles than powders or fresh plants.Usually , the extract ismixe d with the beans asa liquid, and the solvent evaporates before thebean s are stored. Disadvantages that have been mentioned are that extracts aremostl y difficult or laborious to make and yields are usually low.Th e solvents are mostly not available for lowresourc e farmers and large quantities ofplan t material are needed. All plants mentioned inthi s chapter and in table 6wer e used asextract s unless stated otherwise.

Table6: Plants ofwhich extracts are used as protective agentsagainst storage beetles, thesolvent, the extracted plant part, the quantity and the effect. Plant (sub-) Plant species Solvent Plant Quantity ~ Beetle Affected stage # Refe­ A family * part$ species A O E H L M P rence

Acanthaceae Adhatoda vascia B, Et, -- 40 g/kg Cm ** ** ** 111 PE Acanthaceae Andrographis Et W Soaked in Cc *! 117 paniculata 0.125-20%

Annonaceae Annona glabra -- S 0.5% Cm *! 242 Annonaceae Annona muricata -- S 0.5% Cm *! 242 Annonaceae Annona squamosa - s 0.5% Cm *! 242 Annonaceae Monodora myristica EE s Rubbed with Cm H H *! H 210 crude extract Annonaceae Stelechocarpus s 0.5% Cm *! 242 cauliflorus Apiaceae Anethum graveolens Ac s 10-50 Cm, Ls, So, *i 319 Hg/insect Tco Apiaceae Carum roxburghianum PE s 1-5 Cc M 45 Apiaceae Coriandrum sativum Ac s 10-50 Cm, So *i 320 ug/insect

Apocynaceae Nerium oleander PE L 0.1-5%w:v Cc *! Ü M 83 Apocynaceae Rhazya stricta Aq L 0.1% Cm *! *! *! 84 Apocynaceae Strophantus hispidus Et L lOg/l Ao, Pt, So *! 331 Apocynaceae Thevetia peruviana PEB L,T lg/ml, Cc ** ** 230 5-15 ml/1

Arecaceae Cocos nucifera PE Nut 1-3 Cm ** ** ** *i 129 shell Asteraceae Ageratum conyzoides PEB buds, 1 g/ml, Cc ** ** 230 F1.L 5-15 ml/1 Asteraceae Artemisia absinthium Et L 0.5-1.0% Ao ** *! 175

55 6 Extracts

Plant family Plant species Solvent Part Quantity Beetle A O E H L M P Ref. Asteraceae Artemisia dracunculus - ~ 2-8% Ao *! *! 253 Asteraceae Chamomilla recutita - - 2-8% Ao *i 253 Asteraceae Chromolaena odorata Et L 10 g/1 Ao, Pt, So *! 331 Asteraceae Chrysanthemum spp. - Fl 5-100 g/kg Ca *! 139 Asteraceae Eclipta prostrata - ~ 10-50% Cc *! *! 111 Asteraceae Parthenium B,Et, 40 g/kg Cm ** ** »* 111 hysterophorus PE Asteraceae Saussurea lappa PE R,Rh 1-5 Cc H 45 Asteraceae Sphaeranthus indicus PE L,T 0.001-0.05 Cc *! 29 Asteraceae Tagetes minuta Aq R, Fl, L 0-1000 Zs *! 347 Hg/cm2 Asteraceae Tridax precombens B,Et, 40 g/kg Cm *! *! *! 111 PE

Boraginaceae Heliotropium Aq T 0.1% Cm *! *! *! 84 bacciferum

Brassicaceae Lepidium aucheri Aq S 0.1% Cm *! *! *! 84

Capparaceae Boscia senegalensis Ac Fr 1000 g/1, Cm, Pt, Sz, *! 288 5ml Tea

Chenopodiaceae Atriplex lentiformis PE L 0.1-5% w:v Cc *! *! Ü 83

Convolvulaceae Ipomoea carnea PEB L lg/ml, Cc M M 230 5-15 ml/1 B,Et, 40 g/kg Cm ** ** ** 111 PE Convolvulaceae Ipomoea palmata PE L 0.1-5% w:v Cc n *i n 83

Cupressaceae Thujopsis dolabrata Aq Sd 1.5-5 mg Cc, Ls, So *! 7

Cyperaceae Cyperus rotundus Et R 100, Cm *! *! 3 1.25-10%

Euphorbiaceae Ricinus communis Et L 0.5-1.0% Ao *! *! 175 Et S 10 g/1 Ao, Pt, So *! 331 Lamiaceae Hyptis spicigera Aq,E, w lg/ml, Ao *! *! 162 Et, Me 0.3-30 Lamiaceae Hyptis suaveolens Et L 100, Cm ** *! 3 1.25-10% Lamiaceae Mentha piperita - ~ 2-8% Ao *! *! 253 Lamiaceae Ocimum basilicum Es L 2-10 g/kg Cm, So *! 97

56 Theus eo fplan tproduct st oprotec t storedseed s

Plant family Plant species Solvent Part Quantity Beetle A O E H L M P Ref. Lamiaceae Salvia officinalis Aq T 0.1% Cm *! *! *i 84 Lamiaceae Thymus serpyllum -- -- 2-8% Ao *i 253 Lamiaceae Thymus vulgaris - -- 2-8% Ao *i 253

Lecythidaceae Napoleona imperialis PE-Me L 5 g/kg Cm, Csu, Sz ** *# ** 176

Leguminosae- Caesalpinia bonduc PE Fr 1-5 Cc M 45 Caesalpinioideae Leguminosae- Chamaecrista nigricans Aq,E, W lg/ml, Ao *! *! 162 Caesalpinioideae Et, Me 0.3-14 Leguminosae- Erythrophleum Et L 10 g/l Ao, Pt, So *! 331 Caesalpinioideae suaveolens Et Bark 100, Cm *! *! 3 1.25-10% Leguminosae- Senna occidentalis E S (oil) 10 Cm ** ** *! 165 Caesalpinioideae

Leguminosae- Tetrapleura tetraptera Es Fr 2-10 g/kg Cm, So *i 97 Mimosoideae

Leguminosae- Lonchocarpus Et02, S 0.001-0.5% Cm *! 39 Papilionideae salvadorensis CH2C12 Leguminosae- Melilotus officinalis 2-8% Ao *t 253 Papilionideae Leguminosae- Pachyrhizus erosus S 5-100 g/kg Ca *i *! 139 Papilionoideae Leguminosae- Phaseolus vulgaris Aq S peels Bean Cc, Cm ** 108 Papilionoideae equivalent Leguminosae- Pongamia pinnata 2% Cc *! 137 Papilionoideae Leguminosae- Psoralea corylifolia PE S 1-5 Cc H 45 Papilionoideae Leguminosae- Trigonella foenum- Ac, He, L,S 6-30 Ao, Tea *! *! 234 Papilionoideae graecum Me ug/insect, 130 ug/bean Leguminosae- Vignaangularis Aq S peels Bean Ce, Cm ** 108 Papilionoideae equivalent Leguminosae- Vigna unguiculata Aq S peels Bean Ce, Cm ** 108 Papilionoideae equivalent

57 6 Extracts

Plant family Plant species Solvent Part Quantity Beetle A O E H L M P Ref.

Liliaceae Allium cepa PE B 10-50 g/kg Cc ** 228 Et Scale L 100, Cm ** *! 3 1.25-10% PE -- 5% Cm ** 111 Liliaceae Allium sativum PE B 10-50 g/kg Cc *! 228 PE -- 1-3% Cm *! 111 Liliaceae Gloriosa superba -- R 10g/kg Ca ** *» 139

Malvaceae Sida acuta Et L 10 g/1 Ao, Pt, So *! 331

Meliaceae Aglaia elliptica - S 0.5% Cm *! 242 Meliaceae Aphanamixis PE S 10-100 Cc *! 330 polystachya Hg/insect Ac, Et, S 0.123 g/1 Cc, So, Tea *! 329 PE Meliaceae Azadirachta indica Et -- 0.3-30 Ao *! *! 162 -- -- 3-15 Ao *! 237 Et S 10 g/1 Ao, Pt, So *! 331 -- K 10% Ca *i ** ** 100 -- -- 1% Cc *! 137 PEB L, twigs 1 g/ml, 5-15 Cc *! *! 230 Aq K,L 0.1% Cm *! *! *! 84 Aq, Et, S 0.02-1% Cm n 1! M D 169 Me Ac, Aq, L 0.5-3.0% Cm n I) M !! 170 Cf.Et He K 5-200 ^1 Cm M il il ti 259 Aq L,Fr Dipped Cm *! *! *! 287 25-60g/ 1 Aq L,K Soaked in Cm, Cpu, So *! *! *! 172 fatty residue Ac, Aq, FI, L, S 1-3% Cm, Rd, So, *i 275 Et,PE Tea, Tg Meliaceae Dysoxylum cauliflorum -- S 0.5% Cm *! 242 Meliaceae Khaya senegalensis Et W 0.3-30 Ao ** ** 162 Meliaceae Melia azedarach Ac,PE Fr 1,25-10% Cm *! H 85

Menispermaceae Cissampelos owariensis Et L,R 10 g/l Ao, Pt, So *! 331

58 Theus eo fplan tproduct st oprotec t storedseed s

Plant family Plant species Solvent Part Quantity Beetle A O E H L M P Ref.

Myrsinaceae Embelia ribes PE S 1-5 Cc Ü 45 B,Et, 40 g/kg Cm ** ** ** 111 PE

Myrtaceae Eucalyptus citriodora Es L 2-10 g/kg Cm, So *! 97 Myrtaceae Eucalyptus tereticornis Es L 2-10 g/kg Cm, So ** 97 Myrtaceae Eugenia aromatica Et FI 100, Cm *! »! 3 1.25-10% Aq FI 0.1% Cm »! *! »! 84

Piperaceae Lepianthes peltata Et S 100, Cm *! »! 3 1.25-10% Piperaceae Piper guineense Et S 12.5-37.5 Cm *! !! Ü 178 g/kg PE-Me Fr 5 g/kg Cm, Csu, Sz »! »! »! 176 He Fr 5-40 g/1, Cm, Ls, So, »! »! 317 0.5 ni Tco Es Fr 2-10 g/kg Cm, So ** 97 Piperaceae Piper nigrum -- - 0.25-0.5% Cc Ü 111 MC Fr 0.15-20 Cc *! 188 Hg/insect Aq S 0.1% Cm *! *! *! 84 Ac Fr 140 g, Cm *! 321 11.3% yield Ac Corns 50 g/1, 0.5 ni Cm, Ls, So *i H 316 Ac Fr 5-40 g/1, Cm, Ls, So, *i *i 317 0.5 ni Tco Et Fr 3.13-25 Cm, So Ü 315 ug/insect

Poaceae Cymbopogon nardus -- - 3.5% Ao »! »! 327 Poaceae Cymbopogon spp. Et - 0.3-30 Ao *! »! 162

Pontederiaceae Eichhomia crassipes PE L 100-400g/m l Ce Ü 257 Es L 2-10 g/kg Cm, So ** 97 Rubiaceae Coffea arabica Et S Soaked Cc *! »! 266 Rubiaceae Diodia sarmentosa PE-Me FI 5 g/kg Cm, Csu, Sz ** ** ** 176 Rubiaceae Mitracarpus scaber PE-Me FI 5 g/kg Cm, Csu, Sz *• ** ** 176

Rutaceae Citrus limon He P 15-35 Cm, So »! 322 ^g/insect

59 6 Extracts

Plant family Plant species Solvent Part JQuantit y Beetle A O E H L M P Ref. Rutaceae Citrus sinensis Es P 2-10 g/kg Cm, So ** 97 Rutaceae Citrus spp. Aq P 0.1% Cm *! *! *! 84 Rutaceae Zanthoxylum alatum Ac Pericarp 10-50 Cm, Ls, So, *! 318 Hg/insect Tco Rutaceae Zanthoxylum Ac, Aq R 50 Ce *! *! 213 zanthoxyloides

Sapindaceae Dodonaea viscosa PE -- 1-3 Cm ** *! *! H 129

Solanaceae Capsicum annuum Ac Fr 1.25-10%, Cm *! ** 159 200 Solanaceae Capsicum frutescens Ac Fr 1.25-10%, Cm *! *! 159 200 Es Fr 2-10 g/kg Cm, So *! 97 Solanaceae Cestrum nocturnum PE L 1-5 Cc *! 45 Solanaceae Datura metel Et S 10 g/1 Ao, Pt, So *! 331 Solanaceae Datura stramonium Et S 10 g/1 Ao, Pt, So *! 331 Ac, PE, L 5-10% Cm ** *! 195 Et Solanaceae Solanum nigrum Et L 10 g/1 Ao, Pt, So *! 331 Solanaceae Withania somnifera PE R, Rh 1-5 Cc *! 45

Tiliaceae Grewia carpinifolia Et L 10 g/1 Ao, Pt, So *! 331

Verbenaceae Clerodendrum inerme PE L 1-5 Cc *! 45 PE L 0.1-5% w:v Cc *! *! it 83 Verbenaceae Lantana camara PEB L, FI 5-15 g/1 Cc il M 230 Me,PE L,T 1-5% crude Cc *! H 277 -- L 10 Cc, Cm ** *! ** ** *! 245 Es L 2-10 g/kg Cm, So ** 97 Verbenaceae Vitex negundo PE -- 1-3 Cm ** *! ** *! 129

Zingiberaceae Aframomum melegueta Et S 100, Cm ** *! 3 1.25-10% *: Ac = acetone, Aq = aqueous, B= benzene, Cf = chloroform, E = ether, EE = ethyl ether, Es = ester, Et = ethanol, He = hexane, MC = methylene chloride, Me= methanol, PE = petroleum ether, PEB = petroleum ether, diluted with benzene $: B = bulbs, Fl = flowers, Fr = fruits, K = kernels, L= leaves, R = roots, Rh = rhizomes, P =peels , S= seeds, Sd = saw dust, T = twigs, W = whole plant A: See table 1 ~: in ml extract/kg stored product unless stated otherwise #: A= adult longevity & fecundity, O = oviposition, E = survival of eggs & embryos on the seed surface, H = hatching, L= survival of larvae and pupae inside the seed, M = emergence, P= population numbers & effect on the stored product; ** = Measured, but no statistically significant results were found or presented, *!= Significant decrease, !!= Total inhibition.

60 Theus eo fplan tproduct st oprotec tstore dseed s

6.1 DEVELOPMENTAL STAGE AFFECTED

6.1.1 EFFECTS ON ADULTS Extracts often have aneffec t onadults ,eithe r acting asa repellent , atoxicant , ora combination ofthes e two actions.Lantana camera showe d a feeding deterrent action andwa s slightly toxict o Cc,showin g alos so ffecundit y athighe rdose s(277) .Piper guineenseals o caused adult mortality of Cm,Csu an dSz (176 )wherea sPiper nigrum showed contact and oral toxicity against Cm, Ls andSo (316).Fo r Cc, Eichhornia crassipescause dattractio n andmortalit y ofadult s (257),wherea sAphanamixis polystachya was slightly repellent and quitetoxi ct oth ebeetle s (330). Seeds coated with Ocimum basilicum, Piperguineense, Eucalyptus citriodora, Capsicum frutescens and Tetrapleura tetrapteracause d adultmortalit y for Cm(97) .Azadirachta indicaseed s and leaves effectively caused adult mortality of Cm(169 ; 170)an drip ekernel s also reduced fecundity (287),wit hadult sbein gmor e susceptible than eggsan dlarva e(259) . Bosciasenegalensis cause d mortality of Cm, Pt,Sz and Tea (288)wherea sAnethum graveolenseffectivel y repelledSo and Tco(319) .Ricinus communis, Solanumnigrum, Cissampelos owariensis,an dErythrophleum suaveolenswer e alltoxi ct oAo, Pt andSo (331). Trigonellafoenum-graecum reduced longevity and fecundity ofAo and affected the fertility ofbot h sexes of Tea (234).Zanthoxylum zanthoxyloidesdisturbe d mating behaviour of Cm(213) .Th e insecticidal action of Thujopsis dolabrataagains tLs, Cc andSo wa s attributable to fumigant action (7).Root so f Tagetes minutawer e more active against Zstha n flowers and leaves(347) . Single components from extracts can differ inefficac y from thecomplet e extract. Components isolated from Citrus limonshowe d weak toxicity toSo and Cm but were less effective than the complete extract (322).Th e compound, 1,3,7- tritrimethylxanthine, isolated from Coffeaarabica, effectivel y caused sterility (266)an d pipericide, anamid efrom Piper nigrum,wa s highly toxic to Cc(188) .Piperin e inPiper nigrumwa shighl y toxic to adults ofSo (315).Th e complete extract ofPiper guineense (178)an dthre eamide s isolated from Pipernigrum showe d toxicity to Cm(321) , whereasPiper guineense an dPiper nigrum showe d contact toxicity to Cman dSo (317).

6.1.2 EFFECTS ON OVIPOSITION The effects onadult s causeeffect s on oviposition, numbers ofoffsprin g etc.I n some cases,th e eggstha thav ebee n laidar e affected aswell .Lantana camara(277 )an dhig h concentrations ofAndrographis paniculata (117 )reduce d thenumbe r of eggslai db y Cc.Azadirachta indica, Lantanacamara, Ageratumconyzoides, Thevetia peruviana and Ipomoeacarnea, dilute d withbenzen e allrepelle d adult Ccbeetle s and prevented oviposition (230). Azadirachta indicakernel s (170;287) ,Piper guineense (178), Monodoramyristica (210 ) andZanthoxylum zanthoxyloides(213 )inhibite d oviposition by Cm. Lantana camaraha d an effect on oviposition of Ccan d Cm,bu tth e eggstha t werelai d developed normally andth egrain swer e destroyedwit h alittl e retardation only (245).Extract s ofLonchocarpus Salvadorensis, mixed with cowpea flour in artificial seeds,reduce d oviposition of Cm(39) . Capsicumfrutescens was effective in reducing female fecundity andovipositio n of Cm(159) .Dodonaea viscosa (129 )an d Azadirachta indicawer e effective against oviposition, hatching and emergence ofCm (172).Piper guineense wa seffectiv e inpreventin g oviposition of Cm, Csuan dSz (176) . Chamaecrista nigricansan dHyptis spicigera cause d areductio n of oviposition and hatching forAo (162) whereas Trigonellafoenum-graecumonl y inhibited oviposition (234).

61 6 Extracts

6.1.3 EFFECTS ON EGGS AND LARVAE Eggs and young larvae can be affected by extracts whereas older larvae areusuall y less susceptible.Piper guineense cause d eggmortalit y of Cm, Csuan d Sz (176).Capsicum frutescens (159)an dZanthoxylum zanthoxyloides reduce d adult emergence of Cm (213). Monodoramyristica ha d ovicidal and larvicidal effects on Cm(210) ,wherea s eggs were not affected byLonchocarpus salvadorensis, but for this plant, larval mortality was complete and no adults emerged from treated seeds (39).Eggs ,youn g and old larvaeo f Cmwer e susceptible toAzadirachta indica. Eggs failed to hatch, but larvae were more susceptible (259).Azadirachta indica prevented adult emergence (170; 287) and hatching of Cman d was effective against adult emergence ofS o (172).Dodonaea viscosawa s effective against hatching and emergence of Cm(129) , The majority ofth e larvae ofAo were not capable of gnawing orhatchin g after seed treatment withRicinus communisan dArtemisia absinthium. Ricinuscommunis contain s acompoun d that locks the normal metabolism innewl y hatched larvae (175) and hatching was also reduced withHyptis spicigera (162) .

6.2 COMPARISON OFBEETL E SPECIES Not allbeetl e species are equally sensitive toth e extracts.Ao was more sensitive to Ricinuscommunis an dSolanum nigrum tha n Soan dPt were(331) .Whe n applied topically Coriandrum sativum seed was slightly toxic to Tcoan dLs adults,bu t it showed only littletoxi c effect to Soan d itwa stopicall y non-toxic to Cm(320) . Ocimum basilicum, Piperguineense, Eucalyptus citriodora, Capsicum frutescens, and Tetrapleura tetrapterawer etoxi c to Cm,wherea sSo was slightly less sensitive (97). However, So was more susceptible toPiper nigrum than Cm(315 )an d one compound extracted from Citruslemon wa s non-toxic to Cm,bu t slightly toxic toSo (322) . Extracts of different plant parts ofAzadirachta indica ha d aneffec t on oviposition, hatching and emergence for Cm,o n emergence ofS obu t they had no effect onCpu (172).Pericar p of Zanthoxylumalatum showe d no oral toxicity to Cmlarva e orSo adults,ver y little contact toxicity to Cman d Tcoadults ,bu t itwa s topically slightly toxic toSo andLs adults (318).Ls was less sensitive to variousPiper nigrumvarietie s thanSo and Cm were (316).Anethum graveolens wa s non-toxic to Cm,no t verytoxi c to Ls and Tco, but itwa stoxi ct oSo upo ntopica l application (319).Acorus calamuswa s toxict oSo adults,bu t not to Teaadult s (233).Aphanamixis polystachya had a strong repellent effect on Tea andSo, bu t wasno t repellent to Cc.Extract s werehighl y toxict o Ccan d Tea, but only moderately toxic toSo (329) .

6.3 COMPARISON OFPLAN T SPECIES When different plants are compared, striking differences canb e found.Dodonaea viscosawa s effective, but Vitex negundoan d Cocos nuciferadi d notproduc e any effective insecticidal substances against Cm(129) .Ricinus communis an dSolanum nigrumwer e moretoxi ctha nAzadirachta indicaan d the otherplant s tested by Tierto Niber etal. (331) . Capsicumfrutescens was effective inreducin g female fecundity and oviposition of Cm,wherea s Capsicum annuumshowe d no effect (159).Piper nigrum was more effective thanPiper guineense a sa contact toxicant and a grain protective agent against Cm(317) .Piper guineense cause d adult mortality of Cm, Csuan dSz whereasNapoleona imperialis, Mitracarpus scaberan dDiodia sarmentosa didno t show any insecticidal effect (176).

62 Theus eo fplan tproduct st oprotec tstore dseed s

Even ifdifferen t plantpart s or different components ofon e extract are compared, the results can vary greatly.Azadirachta indicafruit s reduced the damage caused by the Fl generation bymor etha n 7time s(285 ) andwer emor e effective thanAzadirachta indica leaves (287). Citruslimon peel s contained four major compounds that were lesstoxi c to adults of Cman dSo than the mixture ofthes e compounds in the complete extract (322). Three amides from Pipernigrum differed inthei r toxicity to adults of Cm.Mal e beetles were more sensitive than females (321).

6.4 DURATION OFTH E EFFECTS The short-term knockdown effect ofplan t extracts canb e important. LD50fo r Cmwa s reached after 24hour s with Ocimum basilicuman d withPiper guineense, wherea s for Eucalyptuscitriodora, Capsicum frutescens, and Tetrapleura tetrapterai ttoo k 48 hours (97). Not much isknow n about the long-term grainprotection . Carum roxburghianuman d Psoraleacorylifolia gav e almost complete protection up to 135days ,an dSaussurea lappa, Withania somniferaan dEmbelia ribes protecte d upt o 90 days(45) .Allium sativumprotecte d stored gram against Ccfo r 135days ,bu t after thisperiod , the damage increased (228).Dodonaea viscosa was effective for lesstha n 33 days only (129).

6.5 EFFECTS ONTH E STORED PRODUCT No effects ofplan t extracts on stored seeds havebee n noted inth e screened literature. Germination of seeds treated withAzadirachta indica remaine d almost equal to control seeds (169; 170).Monodora myristica di d not reduce germination oftreate d cowpea seeds(210) .

6.6 CONCLUSIONS Extracts are usually more effective than powders. They are mostly effective against adult beetles, either asrepellent s or astoxicants .Othe r developmental stages ofth e beetle can be susceptible aswell . The effect ofextract s onth e stored product isusuall y negligible. Large quantities ofplant s are often needed to obtain sufficient amounts of extracts. Solvent availability can imposea limitation onth eus eo fextracts .

63 The use ofplan t products toprotec t stored seeds

7GENERA L DISCUSSION For each plant species, sorted alphabetically inTabl e 7,th e common name,th e descriptor and a summary of the information from literature, with regard to the different types of applications for thecontro l of seedbeetle s are given. Inthi s table, several plants are mentioned for which the use isnote d as unknown. In the cited references, these plants have been mentioned, often astraditionall y used or effective plants,withou t enough detail about their application tojustif y incorporation inth e separate tables in earlier chapters. Relevant synonyms from the scientific nomenclature are shown in the tablewit h areferenc e toth enam etha tha sbee nuse d inth e other tables inthi s review.

7.1 MODEO F ACTION The authors of thepublication s cited in this review tried to assess whether plants are effective in protecting stored seeds from beetle attack. Some focussed on adult beetles, others on larvae or onth e insect population as awhole . The effect of thetreatmen t and the most affected developmental stage can be found only ifth ebeetle s are examined during their whole developmental cycle,whic h israrel y done.Fo ruser s ofplan t material against insect pests and for many ofth e cited authors,th e eventual effect is usually more important than the mechanism of action against each developmental stage ofth e beetles. However, knowledge ofth emechanis m mayb e essential inth e development of effective applications for beetle control. The articles mostly refer to laboratory tests with aplan t product applied to uninfested beans as loose grains to which beetles are added. However, traditionally, beans are often infested at harvest and stored in their pods.Efficac y ofpowders , ashes,volatil e oilsan d extracts could be different when used under these conditions.Th epod s around the beans would serve asa n extra barrier for the beetle but atth e same time,the y would act asa barrier between the seeds and the protective plant product. Application ofnon-volatil e oils isonl y effective if looseseed sar e stored. Thepo d with its cracks,pubescenc e and rough surface would not be appropriate to apply afilm o foi l to.

7.2 COMPARISON OF BEETLE SPECIES Arang e ofbeetl e species were subjected tobioassays , although most ofth e studied species belong to the families ofBruchida e and Curculionidae. Some authors refer to comparative research between two ormor eo fthes e beetle species or families, but the range of species compared in such publications is rarely the same.Non e of the tested beetle species has been analysed thoroughly enough to know all itsproperties . Therefore, none of thebeetl e species could serve asa contro l species with known or predictable responses to seed treatments. Plant products can therefore not be ranked for their toxicity according tothei r effect on such abeetl e species. The reactions ofdifferen t biotypes or of individual beetles within a species canb e variable. Beetles ofth e same species,bu t ofdifferen t geographical origin have been found to show differences in life history and inrespons e totreatments , even after their rearing inth e laboratory for several generations (70; 56).Difference s between females and males,whic h often differ in size,hav ebee n reported as well.

65 7 General discussion

7.3 COMPARISON OF APPLICATION METHODS The used plant materials are inherently biologically variable and can show considerable differences in activity. Forplants ,th e composition and contents of secondary metabolites, usually involved in defence ofth eplant , can differ greatly between ecotypes,an d even between individuals. The amount of these components perplan t can vary with, among others,th e growth and storage conditions, with the time of harvest and with the plant part. Forplan t extracts, the results can also be dependent on the quality and purity of the solvents. The applied dosages are often expressed in different units.Unit s of weight orvolum e are not uniformly used. Furthermore, in many studies the plant material was applied to filter paper ordirectl y to various developmental stages of the insects, not to the seeds. In some cases plant materials are as effective as,o r even more effective than synthetic insecticides.Zanthoxylum zanthoxyloides(212 ) andAzadirachta indicawer e as effective aspirimiphos-methy l and permethrin against Cm(211) . Tamarindus indicaas h was as effective as actellic (pirimiphos-methyl) dust against Tg(5) .N o significant difference in effect against bruchids was found between Zeamays oil ,pirimiphos - methyl, and permethrin dust (309). Striking differences exist between approaches of application methods ofplan t material to stored seeds.Fo r non-volatile oils,ther e are only few plants tested, but for each oil many references can be found. Forpowder s and extracts,th e opposite istrue , many plant species have been tested, but perplan t only few references are found. Different applications of the sameplan t will show different effects. Secondary products such as oils and extracts are usually more effective than powders or whole plant parts from which they were obtained. Seed powder ofPiper guineense was less effective than its ether extract in enhancing adult mortality in Cm(178) . Oil of Azadirachta indicaseed s was much more effective than the seed powder. This was mainly because the oil could penetrate into the egg whereas powders cannot do so (161). Treatment ofbean s with oil even at less than 1 ml/kg was more effective than treatment with ashes (338). Depending on the quantity used and the duration ofth e experiment, the efficacies of volatile and non-volatile oils can vary. Volatile oil ofPiper guineense was more effective than non-volatile oil ofAzadirachta indica, agains t Cm(124) .Oil s of Jatropha curcas, Azadirachtaindica, Calophyllum inophyllum andMadhuca longifoliawer e all effective against Cm,wherea sth e volatile oil ofEugenia aromatica was much less effective (129). Ifplan t products are mixed, again different activities can be found. Efficacy can be enhanced or decreased. For instance, powder ofPiper nigrumgav e abette r protection of stored seeds when used in conjunction with non-volatilemustar d oil (149).Ver y few studies explored thepossibilit y ofusin g mixtures. Potentially the use oftw o or moreplan t products could enhance their efficacy, especially when the different components act along different mechanisms. Care should be taken that the effect measured is really attributable to the plant and that other factors are stable and ineffective. Azadirachtaindica, Lantanacamara, Ageratumconyzoides, Thevetia peruviana andIpomoea carnea, dilute d with benzene all repelled adult Ccbeetle s and prevented oviposition. However, pure benzene had the same effect asth e extracts (230).

66 Theus eo fplan tproduct st oprotec tstore dseed s

7.4 DURATION OF THE EFFECTS Ifth eperio d of action ismentione d in the literature, it iseithe r presumed to be the timeneede d tokil l theadul t beetles orth eperio d over which theproduc t provides satisfactory protection against the insects. In some cases,th e time ismeasure d in which apopulatio n of acertai n number ofbeetle s can build up, or in which acertai n percentage ofdamag e is inflicted. Efficacy of plant products is shown by either their postponing effect onth e development of individual beetles or of abeetl e population, or as adecreas e inth e damage after acertai n amount of time. Long-term (more than awee k of incubation) and short-term (a few hours to aweek ) tests each makeu pabou t half ofth e total number of experiments in the screened articles.

7.5 EFFECT ON THE STORED PRODUCT Beans are affected by storage even ifinsec t infestation does not occur. Cooking time, seed germination and seedling vigour can be affected. Forthos e reports whereth e treated samplesperfor m bettertha nth econtrol ,i t isno t alwaysclea r whether the control sample has been infested with beetles ornot . Seeds, from which adult beetles have emerged, do not germinate readily. If theeffec t ofth e treatment with (effective) plant products ist o be analysed, this should be done against seeds that are asmuc h infested as the treated seeds,otherwis e the effect ofbruchi d infestation on seed properties wouldb e tested. Many ofth e effects, such as changes ofcolou r and effects ontast e of the seeds duet o decomposition of theapplie d plant material, are only important ifth e seeds aret ob e sold or eaten. For sowing seeds, only germination and seedling vigour need tob e preserved or improved by the treatment.

7.6 CONCLUSION Seed beetles cause an important part ofth etota l insect damage to seed crops.I nstore , they usually are the main pests. Much research has already been done to develop methods to prevent this damage. Many different plant species have been tested for their effect on the beetles. However, no general ranking of insecticidal or repellent effect can be given. This lack ofrankin g is on the onehan d due to the lack of experimental detail and statistical analyses ofth eresult s in many of the cited publications. Onth e other hand,th epotentia l efficacy of aplan t product can depend onth eplan t species,th e individual plant, the plant part, and the time and way ofharvesting . In bioassays,th e application mode,th e quantity, the preparation ofth e plant material, the state (humidity, intactness ofth e seed skin and/orpod ,rat e of infestation) ofth eproduc t to store andth e tested beetle species are important for the outcome ofth etests .Th e mere fact that natural products areused , impliestha tconsiderabl e variation ist ob e expected inth e outcome of the experiments. Results in literature are often contradictory, but many ofth eteste d plants dosho w effects against seed beetles.Th e most effective plants or methods of application areno t yet known, but results are promising and plant material canb e aneffectiv e weapon in the battle against the beetles. Plants canb e an effective replacement for chemical insecticides toprotec t stored seeds.

67 7 General discussion

Table7: Common namesof the plants and the type of application reported Plant species Author Common name Plant (sub-) Application family A E O P V Unknown 1 Acacia albida Del. Winter thorn Leguminosae 34; 35; 113 Mimosoideae Acacia arabica Willd. SeeAcacia nilotica Leguminosae Mimosoideae 2 Acacia concinna DC. Soap pod Leguminosae 1 109 Mimosoideae 3 Acacia laeta L. Leguminosae 113 Mimosoideae 4 Acacia nilotica (L.) Willd. ex Delile Gum Arabic tree/ Leguminosae 1 109 babul acacia Mimosoideae 5 Acacia Senegal (L.) Willd. Gum arabic Leguminosae 34; 35; 114 Mimosoideae 6 Acacia sinuata (Lour.) Merr. Soapnut Leguminosae 2 Mimosoideae 7 Acacia spp. Mill. Acacia Leguminosae 1 65 Mimosoideae 8 Achyranthes aspera L. Devil's horsewhip Amaranthaceae 1 9 Acorus calamus L. Sweetflag Araceae 5 8 65; 68; 95; 109; 164; 224; 233; 258; 312; 336; 351 10 Adhatoda spp. Mill. Water willow Acanthaceae 1 11 Adhatoda vascia Nees Malabar nut Acanthaceae 1 2 65; 95; 340 12 Aerva javanica (Burm. f.) Juss. ex Java aerva Amaranthaceae 15; 18; Schult. 114; 171 13 Aframomum Schum. Melegueta pepper Zingiberaceae 1 1 melegueta 14 Afrormosia laxiflora (Benth. ex. Bak.) Leguminosae- 103; 109 Harms Papilionoideae 15 Afzelia africana Sm. ex Pers. African mahogany Leguminosae- 3 114 Caesalpinioideae 16 Ageratum conyzoides L. Goat weed/ Asteraceae 1 1 1 95; 198 tropical whiteweed 17 Ageratum Mill. Bluemink Asteraceae 109 houstonianum 18 Aglaia elliptica Blume Meliaceae 1

68 The use of plant products to protect stored seeds

Plant species Author Common name Plant family A E O P V Unknown 19 Ajuga remota Benth. Bugle Lamiaceae 109 20 Allium cepa L. Garden onion Liliaceae 3 95; 208; 312; 336; 340 21 Allium sativum L. Cultivated garlic Liliaceae 2 2 65; 109; 312; 340 22 Alnus nepalensis D.Don Nepal alder Betulaceae 109 23 Aloe marlothii A. Berger Mountain aloe Liliaceae 1 24 Alpinia galanga (L.) Sweet Greater galangal/ Zingiberaceae 2 62 ginger lily 25 Alpinia zerumbet (Pers.) Burtt & R.M. Shell ginger Zingiberaceae 62 Sm. 26 Alysicarpus (Schumach.) J. Alyce clover Leguminosae- 34; 35 ovalifolius Léonard Papilionoideae 27 Amaranthus L. Pigweed Amaranthaceae 15; 18; graecizans 114; 171 Amoora rohituka Wight & Arn. See Aphanamixis Meliaceae polystachya 28 Anacardium L. Cashew Anacardiaceae 2 65; 109 occidentale 29 Ancistrocladus Scott-Elliot Ancistroclada- 116 barteri ceae 30 Andrographis (Burm. f.) Wall, ex False waterwillow Acanthaceae 1 paniculata Nees Andropogon citratus DC. ex Nees See Cymbopogon Poaceae citratus 31 Anethum graveolens L. Dill Apiaceae 1 1 1 336 Annona arenaria Thonn. See Annona Annonaceae senegalensis 32 Annona glabra L. Pond apple Annonaceae 1 33 Annona muricata L. Soursop Annonaceae 1 312 34 Annona reticulata L. , Custard apple Annonaceae 1 65; 109; 249; 252; 312 35 Annona senegalensis Pers. Wild custard apple Annonaceae 2 18; 19; 34; 35; 103; 109; 114; 171; 270 36 Annona spp. L. Annona Annonaceae 1

69 7 General discussion

Plant species Author Common name Plant family A E O P V Unknown 37 Annona squamosa L. Sweetsop/ Annonaceae 1 3 109; 312; sugar apple 336 38 Anogeissus (DC.) Guill. & Combretaceae 34;35; 113 leiocarpus Perr. 39 Aphanamixis Wall. & Parker Pithraj Meliaceae 2 1 184 polystachya 40 Apium graveolens L. Wild celery Apiaceae 1 41 Apium spp. L. Celery Apiaceae 291 42 Arachis hypogaea L. Groundnut/peanut Leguminosae- 48 34; 35; 65; Papilionoideae 66; 95; 109; 114; 164; 258; 312; 336; 339 43 Argemone mexicana L. Mexican prickly Papaveraceae 109 poppy 44 Artemisia absinthium L. Absinth sagewort Asteraceae 1 45 Artemisia L. Green sagewort Asteraceae 1 120 dracunculus 46 Artemisia spp. L. Wormwood/ Asteraceae 65; 109; sagewort 291 47 Artemisia tridentata Nutt. Big sagebrush Asteraceae 1 48 Aspilia afhcana (Pers.) C. D. Adams Asteraceae 1 49 Atriplex lentiformis (Torr.) S. Watson Quail bush/ Chenopodiaceae 1 95 salt bush 50 Azadirachta indica Juss. Neem Meliaceae 2 13 34 35 13; 18; 19; 23; 24; 26; 27; 34; 35; 65; 66; 68; 94; 95; 109; 114; 119; 164; 171; 173; 184; 198; 205; 247; 252; 258; 270; 274; 281; 312; 328; 336

70 The use of plant products to protect stored seeds

Plant species Author Common name Plant family A E O P V Unknown 51 Balanites aegyptiaca (L.) Delile Desert date Zygophyllaceae 1 34; 35; 114;116 52 Balsamorhiza (Pursh) Nutt. Arrowleaf Asteraceae 1 sagittata balsamroot 53 Barringtonia (L.) Gaertn. Freshwater Lecythidaceae 139 acutangula mangrove 54 Barringtonia asiatica (L.) Kurz Lecythidaceae 312 Bassia latifolia Roxb. See Madhuca Sapotaceae longifolia 55 Blumea balsamifera (L.) DC. Sambong/ Asteraceae 1 198 nagi camphor 56 Boesenbergia Schi. Fingerroot Zingiberaceae 62 padurata 57 Borassus aethiopum C. Mart. Fan, palmyra palm Arecaceae 34; 35; 113; 114 58 Boscia salicifolia Oliv. Willow shepherds Capparaceae 15; 18; 34; tree 35; 114 59 Boscia senegalensis (Pers.) Lam. ex Senegal boscia Capparaceae 1 7 18; 19; 24; Poir. 34; 35; 66; 109; 114; 171; 270 60 Bougainvillea spp. Comm. ex Juss. Bougainvillea Nyctaginaceae 109 61 Brassica juncea (L.) Czem. Rape/ Brassicaceae 9 95; 109; Indian mustard 336 62 Brassica râpa L. Turnip/ Brassicaceae 109 rape mustard 63 Brassica spp. L. Mustard Brassicaceae 24 65; 109; 149; 312; 336 64 Bridelia ferruginea Benth. Euphorbiaceae 1 Butyrospermum (Gaertn. f.) Hepper See Vitellaria Sapotaceae paradoxum paradoxa Butyrospermum (G. Don) Kotschy See Vitellaria Sapotaceae parkii paradoxa 65 Cactus spp. Cactus Cactaceae 65; 109 66 Cadaba farinosa Forssk. Cadaba Capparaceae 34; 35 67 Caesalpinia bonduc (L.) Roxb. Yellow nicker/ Leguminosae- 1 bonduc nut Caesalpinioideae

71 7 General discussion

Plant species Author Common name Plant family A E O P V Unknown 68 Caesalpinia (L.) Sw. Dwarf poiciana Leguminosae- 109 pulcherrima Caesalpinioideae 69 Caesalpinia sappan L. Sappan wood Leguminosae- 139 Caesalpinioideae 70 Callistemon citrinus (Curtis) Stapf Crimson bottlebrush Myrtaceae 1 186 Callistemon (Sw.) DC. See Callistemon Myrtaceae lanceolatus citrinus 71 Callistephus (L.) Nees China aster Asteraceae 109 chinensis 72 Calophyllum L. Alexandrian laurel/ Clusiaceae 3 336 inophyllum undi 73 Calotropis gigantea (L.) Aiton f Giant milkweed Asclepiadaceae 109 74 Calotropis procera (Ait.) W.T. Aiton Roostertree/ Asclepiadaceae 15; 18; 34; swallowwort 35; 114; 171 75 Canna indica L. Indian shot Cannaceae 109 76 Cannabis saliva L. Marijuana Cannabinaceae 109 77 Capsicum annuum L. Chilli, hot, cayenne Solanaceae 1 5 34; 35; 95; pepper 151; 208; 252; 336 78 Capsicum chinense Jacq. Aji Solanaceae 1 79 Capsicum frutescens L. Piment/ Solanaceae 2 8 24; 65; 66; red pepper 198; 284; 312; 339 80 Capsicum spp. L. Pepper Solanaceae 1 15; 18; 19; 35; 86; 109; 114; 171; 312; 336 81 Carica papaya L. Papaya/ Caricaceae 258; 270; pawpaw 284 82 Carthamus tinctorius L. Safflower Asteraceae 12 65; 109; 111; 312 83 Carum Benth. ex Kurz Bishops weed Apiaceae 1 291 roxburghianum Caryophyllus L. See Eugenia Myrtaceae aromaticus aromatica

72 The use of plant products to protect stored seeds

Plant species Author Common name Plant family A E JO P V Unknown Cassia nigricans Vahl See Chamaecrista Leguminosae- nigricans Caesalpinioideae Cassia occidentalis L. See Senna Leguminosae- occidentalis Caesalpinioideae Coffee sena 84 Cassia singueana Del. Cassia tree Leguminosae- 15; 18; 34; Caesalpinioideae 35; 114; 171; 270 85 Cassia spp. L. Cassia Leguminosae- 109 Caesalpinioideae 86 Castanospermurn Cunn. Black bean/ Leguminosae- 328 australe moreton bay Papilionoideae chestnut 87 Casuarina indica Pers. Casuarinaceae 1 88 Casuarina spp. L. Casuarina Casuarinaceae 65; 109 89 Cedrus deodara (Roxb. ex D. Don) Deodar cedar Pinaceae 1 G. Don f. 90 Cedrus spp. Trew Cedar Pinaceae 1 91 Ceiba pentandra (L.) Gaertn. Kapok tree Bombacaceae 1 2 92 Celastrus angulata Maxim. Bittersweet Celastraceae 109 93 Centaurea perrottetii DC. Asteraceae 34; 35 94 Cestrum nocturnum L. Night flowering Solanaceae 1 jasmine 95 Chamaecrista (Vahl) Greene Leguminosae 1 6 23; 65; nigricans Caesalpinioideae 114; 153; 312; 336; 339 96 Chamomilla recutita (L.) Rauschen German chamomile Asteraceae 1 120; 291 97 Chenopodium L. Mexican tea Chenopodiaceae 1 95 ambrosioides 98 Chromolaena (L.) R.M. King & Siam weed/ Asteraceae 1 4 2 odorata H. Rob. hagonoy 99 Chrozophora Vis. Chrozophora Euphorbiaceae 34; 35 brocchiana 100 Chrysanthemum (Trevir.)Vis . Pyrethrum Asteraceae 312 cinerariifolium 101 Chrysanthemum L. Manzanilla Asteraceae 1 198 indicum 102 Chrysanthemum spp. L. Daisy Asteraceae 1

73 7 General discussion

Plant species Author Common name Plant family A E O P V Unknown 103 Cinnamomum (L.) J. Presl Camphor tree Lauraceae 2 camphora 104 Cinnamomum Vid. Cinnamon Lauraceae 1 mercadoi 105 Cinnamomum spp. Schaeff. Cinnamon Lauraceae 65 106 Cinnamomum verum J. Presl Cinnamon Lauraceae 1 109;333 Cinnamomum Blume See Cinnamomum Lauraceae zeylanicum verum 107 Cissampelos P. Beauv. ex DC. Pareira brava/ Menispermaceae 1 owariensis velvet leaf 108 Cissus L. Pirandai/ Vitaceae 153 quadrangularis petch-sang-kart 109 Citrullus colocynthis (L.) Schrad. Water melon/ Cucurbitaceae 34; 35 colocynth 110 Citrus aurantifolia (Christm.) Swingle Lime Rutaceae 2 5 65; 109 Citrus aurantium L. See Citrus sinensis Rutaceae 111 Citrus crematifolia Lush. Rutaceae 2 112 Citrus grandis (L.) Osbeck Pomelo Rutaceae 1 113 Citrus limon (L.) Burm. f. Lemon Rutaceae 1 3 7 109; 252; 291;336 114 Citrus parodist Macfad. Grapefruit Rutaceae 3 5 109; 291 115 Citrus reticulata Blanco Tangerine/ Rutaceae 2 4 mandarin 116 Citrus sinensis (L.) Osbeck (sweet) orange Rutaceae 1 8 5 109; 291 117 Citrus spp. L. Citrus Rutaceae 1 1 65; 109; 164;336 118 CitrusX tangelo J.W. Ingram & H.E. Tangelo Rutaceae 2 Moore 119 Clematis vitalba L. Travellers joy/ Ranunculaceae 65; 109 evergreen clematis 120 Clerodendrum (L.) Gaertn. Garden quinine/ Verbenaceae 2 86; 95 inerme embrert 121 Cocculus trilobus (Thunb.) DC. Huehue Menispermaceae 109 122 Cocos nucifera L. Coconut palm Arecaceae 1 33 65; 95; 109; 164; 198; 247; 312; 336 123 Coffea arabica L. Arabian coffee Rubiaceae 1 1

74 The use ofplan t products toprotec t stored seeds

Plant species Author Common name Plant family A E O P V Unknown 124 Coleus amboinicus Lour. Oregano Lamiaceae 1 198; 291; 292; 293 125 Combretum Sond. Red bushwillow Combretaceae 1 apiculatum 126 Combretum imberbe Wawra Motswere tree/ Combretaceae 3 1 leadwood 127 Conomorpha DC. Myrsinaceae 109; 116 peruviana 128 Coriandrum sativum L. Coriander/ Apiaceae 1 2 291 Chinese parsley 129 Crinum defixum Ker. Gawl. Swamplily/ Amaryllidaceae 1 kesarachettu 130 Crotalariajun cea L. Sun hemp Leguminosae- 65; 109 Papilionoideae 131 Crotalaria G. Don. Crotalaria/sunhemp/ Leguminosae- 312 ochroleuca slender leaf Papilionoideae rattlebox 132 Croton Baill. Bonpland's croton Euphorbiaceae 168 bonplandianus 133 Croton gratissimus Burch. Croton Euphorbiaceae 1 134 Croton tiglium L. Purging croton Euphorbiaceae 312 135 Cucurbita Kunth Missouri, buffalo Cucurbitaceae 116 foetidissima gourd 136 Cucurbita spp. L. Gourd/cucurbit Cucurbitaceae 109 137 Cuminum cyminum L. Cumin Apiaceae 1 138 Curcuma aeruginosa Roxb. Black temu Zingiberaceae 62 139 Curcuma amada Roxb. Mango ginger Zingiberaceae 1 62 140 Curcuma aromatica Salisb. Wild turmeric Zingiberaceae 62 Curcuma domestica Val. See Curcuma longa Zingiberaceae 141 Curcuma heyeana Zingiberaceae 62 142 Curcuma longa L. Common turmeric Zingiberaceae 3 65; 66; 312; 340 143 Curcuma zedoaria (Christm.) Roscoe Zedoary Zingiberaceae 1 62 144 Cymbopogon citratus (DC. ex Nees) Stapf Lemon grass Poaceae 5 8 111; 291 145 Cymbopogon (Höchst.) Chiov. Poaceae 34; 35; 114 giganteus 146 Cymbopogon nardus (L.) Rendle Citronella grass Poaceae 1 1 1 7 111

75 7 General discussion

Plant species Author Common name Plant family A E O P V Unknown 147 Cymbopogon (L.) Spreng. Camel grass Poaceae 2 3 66; 114 schoenanthus 148 Cymbopogon spp. Spreng. Cymbopogon Poaceae 1 65; 109 149 Cyperus rotundus L. Nutgrass Cyperaceae 1 Datura fastuosa L. See Datura metel Solanaceae Datura innoxia Mill. See Datura metel Solanaceae 150 Datura metel L. Thornapple/ Solanaceae 1 34; 35 angel trumpet/ horn of plenty 151 Datura stramonium L. Jimson weed/ Solanaceae 2 65; 103; thorn apple 109; 167; 284 152 Delonix regia (Bojer ex Hook.) Royal poinciana Leguminosae- 109 Raf. Papilionoideae 153 Dennettia tripetala Baker f. Dennettia Annonaceae 1 1 65;312 154 Derris elliptica (Wall.) Benth. Jewelvine/derris Leguminosae- 65; 66; Papilionoideae 109; 139; 312 155 Derris inudata Lour. Derris Leguminosae- 1 Papilionoideae 156 Derris malaccensis Prain/Hend. Leguminosae- 312 Papilionoideae 157 Derris spp. Lour. Derris Leguminosae- 1 Papilionoideae 158 Derris trifoliata Lour. Threeleaf derris Leguminosae- 312 Papilionoideae Derris uliginosa (Willd.) Benth. SeeDerris trifoliata Leguminosae- Papilionoideae 159 Diheteropogon Hitchc. Poaceae 113 hagerupii 160 Dillenia retusa Thunb. Dilleniaceae 1 249 161 Diodia sarmentosa Sw. Tropical Rubiaceae 1 1 buttonweed 162 Diplolophium Turcz. Apiaceae 3 africanum 163 Dodonaea viscosa (L.) Jacq. Florida hopbush/ Sapindaceae 1 dodonaea 164 Dolichos buchananii Harms Leguminosae- 116 Papilionoideae

76 The use ofplan t products to protect stored seeds

Plant species Author Common name Plant family A E O P V Unknown 165 Dolichos Taub. Lablab Leguminosae- 109 kilimandscharicum Papilionoideae 166 Duranta plumieri L. Duranta Verbenaceae 86 167 Dysoxylum Hiern Meliaceae 1 cauliflorum 168 Echinochloa Link Japanese millet Poaceae 109 frumentacea 169 Eclipta prostrata (L.) L. False daisy Asteraceae 1 170 Eichhornia crassipes (Mart.) Solms Water hyacinth Pontederiaceae 2 171 Elaeis guineensis Jacq. African oil palm Arecaceae 23 65; 95; 109; 258; 336; 339 172 Eleusine coracana L. Gaertn. Finger millet/ragi Poaceae 65; 109 173 Embelia ribes Brun Baibarang Myrsinaceae 2 95 174 Emilia sonchifolia (L.) DC. Lilac tasselflower Asteraceae 1 175 Entada africana Guill. & Perr. Leguminosae 1 Mimosoideae 176 Eragrostis tremula Höchst, ex Steud. Lovegrass Poaceae 15; 18; 114; 171 Eruca saliva Mill. See Eruca vesicaria Brassicaceae 177 Eruca vesicaria (L.)Cav. Rocket salad Brassicaceae i 2 Eiythrophleum G. Don See Erythrophleum Leguminosae- guineense suaveolens Caesalpinioideae 178 Eiythrophleum (Guill. & Perr.) Sasswood Leguminosae- 2 1 103 suaveolens Brenan Caesalpinioideae 179 Eucalyptus Hook. Lemon scented Myrtaceae 1 1 4 164;223 citriodora eucalyptus 1S0\Eucalyptus globulus Labill. Fever tree/ Myrtaceae 1 1 68; 86; Tasmanian blue 164;223 gum 181 Eucalyptus spp. L'Hér. Eucalyptus Myrtaceae 4 1 65; 91; 109;312

\82\Eucalypttts Sm. Forest red gum Myrtaceae 1 1 Itereticorms \83\Eugenia aromatica (L.) Baill. Cloves Myrtaceae ; i2 1 3 358 Eugenia Thunb. See Eugenia Myrtaceae [ Icaryophyllata aromatica i 184 Eugenia uniflora L. Surinam cherry Myrtaceae j1 |2

77 7 General discussion

Plant species Author Common name Plant family A E O P V Unknown 185 Eupatorium (Lam.) Small Dogfennel Asteraceae 1 186 capillifolium Eupatorium L. See Chromolaena Asteraceae odoratum odorata 186 Eupatorium spp. L. Thoroughwort/ Asteraceae 1 291 fennel 187 Euphorbia Aiton Balsam spurge Euphorbiaceae 15; 18; 34; bahamifera 35; 114; 171 188 Euphorbia poissonii Pax Tinya Euphorbiaceae 34; 35 189 Euphorbia tirucalli L. Milk bush/ Euphorbiaceae 1 Indian tree spurge 190 Feretia apodanthera Del. Rubiaceae 34; 35 191 Ficus exasperata Vahl Moraceae 1 192 Ficus sycomorus L. Sycamore, common Moraceae 18; 114; cluster fig 171 193 Fortunella spp. Swingle Kumquat/fortunella Rutaceae 2 194 Galega spp. L. Galega/rue Leguminosae- 1 291 Papilionoideae 195 Geranium Fisch. & CA. Mey. Sticky geranium Geraniaceae 1 viscosissimum ex CA. Mey. 196 Gloriosa superba L. Flame lily Liliaceae 1 197 Glycine max (L.) Merr. Soybean Leguminosae- 22 65; 91; 95; Papilionoideae 109; 173; 293; 312; 336 198 Gossypium hirsutum L. Upland cotton Malvaceae 17 65; 109; 114; 171; 194; 293; 312; 336 199 Grewia carpinifolia Juss. Tiliaceae 1 200 Guiera senegalensis J. F. Gmel. Combretaceae 19; 34; 35 201 Guizotia abyssinica (L. f.) Cass. Ramtilla/niger Asteraceae 2 95 202 Haplophyton A. DC Cockroach plant Apocynaceae 312 cimicidum 203 Harpullia arborea (Blanco) Radlk. Uas Sapindaceae 116 204 Harrisonia Oliv. Simaroubaceae 109 abyssinica

78 The use of plant products to protect stored seeds

Plant species Author Common name Plant family A E O P V Unknown 205 Helianthus annuus L. Common sunflower Asteraceae 16 65; 109; 173; 258; 312 206 Heliotropium Forssk. Boraginaceae 1 bacciferum 207 Hibiscus chinensis L. Malvaceae 109 208 Hibiscus sabdariffa L. Roselle Malvaceae 15; 18; 34; 35; 114; 171 209 Hyptis spicigera Lam. Marubio/ Lamiaceae 1 5 1 13; 20; 23; black sesame 34; 35; 65; 66; 95; 103; 109; 114; 180; 312; 336; 339 210 Hyptis suaveolens (L.) Poit. English basil/pignut Lamiaceae 1 2 2 34; 35; 180 211 Imperala cylindrica (L.) Beauv. Ekon grass/ Poaceae 109 cogongrass 212 Inula graveolens (L.) Desf. Dill Asteraceae 65; 109 213 Ipomoea asarifolia (Desr.) Roem. & Convolvulaceae 34; 35 Schult. 214 Ipomoea carnea Jacq. Gloria de la manana Convolvulaceae 2 2 95 215 Ipomoea mauritiana Jacq . Convolvulaceae 1 216 Ipomoea palmata Forssk. Convolvulaceae 1 95 217 Jacquinia barbasco Mez Braceletwood Theophrastaceae 116 218 Jasminum spp. L. Jasmine Oleaceae 109 219 Jatropha curcas L. Physic, Barbados Euphorbiaceae 1 312; 336 nut 220 Jatropha indica Jatropha Euphorbiaceae 1 1 221 Juglans spp. L. Walnut Juglandaceae 1 222 Juslicia adhatoda L. , Malabar nut tree Acanthaceae 109 223 Kaempferia galanga L. Galanga Zingiberaceae 62; 109 224 Kaempferia rotunda L. Round rooted Zingiberaceae 62 galangal 225 Khaya senegalensis j(Desr.) A. Juss. African mahogany Meliaceae 1 2 23; 24; 34; 35; 65; 109 226 Lannea fruticosa Engl. Anacardiaceae 15; 18; 114; 171

79 7 General discussion

Plant species Author Common name Plant family A E O P V Unknown 227 Lantana camara L. Aripple/ Verbenaceae 4 4 2 95 common lantana 228 Lantana rugosa Thunb. Verbenaceae 65; 103; 109 229 Lantana salviifolia Verbenaceae 109 230 Lantana spp. L. Lantana Verbenaceae 109 231 Laurus nobilis L. Sweet laurel, bay Lauraceae 2 2 291; 292; leaves 293 232 Lavandula Mill. English lavender Lamiaceae 1 2 292; 293 angustifolia Lavandula officinalis Chaix See Lavandula Lamiaceae angustifolia 233 Lavandula spp. L. Lavender Lamiaceae 2 291 234 Ledum palustre L. Marsh Labrador tea Ericaceae 1 235 Lepianthes peltata (L.) Raf. Monkey's hand Piperaceae 1 236 Lepidium aucheri Boiss. Aucher's Brassicaceae 1 pepperwort 237 Leptadenia hastata (Pers.) Decne. Spear leaved Asclepiadaceae 19; 34; 35 leptadenia 238 Linum usitatissimum L. Linseed/ Linaceae 6 95 common flax 239 Lippia adoensis Höchst, ex Walp. Verbenaceae 1 2 240 Lippia chevalieri Moldenke Verbenaceae 1 66 241 Lippia multiflora Moldenke Verbenaceae 1 4 35 242 Lobelia columnaris Campanulaceae 284 243 Lobelia nicotianifolia Heyne Wild tobacco Campanulaceae 109 244 Lonchocarpus Pittier Leguminosae- 1 salvadorensis Papilionoideae 245 Lonchocarpus spp. Kunth Lonchocarpus/ Leguminosae- 66 lancepod Papilionoideae 246 Luffa acutangula (L.) Roxb. Sinkwa Cucurbitaceae 109; 116 towelsponge/ dishcloth gourd 247 Luffa aegyptiaca Mill. Sponge gourd Cucurbitaceae 103; 109 Luffa cylindrica (L.) M. Roem. See Luffa Cucurbitaceae aegyptiaca 248 Luvunga scandens Buch Ham. Sugandh kokila Rutaceae 71

80 Theus eo fplan tproduct st oprotec tstore dseed s

Plant species Author Common name Plant family A E O P V Unknown 249 Lysimachia L. Moneywort/ Primulaceae 65; 109 nummularia creepingjenn y Madhuca indica J.F. Gmel. See Madhuca Sapotaceae longifolia Madhuca latifolia (Roxb.) J.F. Macbr. See Madhuca Sapotaceae longifolia 250 Madhuca longifolia (J. König) J.F. Mahua/ Sapotaceae 7 65; 95; Macbr. illipe butter tree 109; 111; 336 251 Maerua angolensis DC. Bead bean Capparaceae 153 252 Maerua crassifolia Forssk. Atil/toothbrush tree Capparaceae 15; 18; 34; 35 253 Mammea americana L. Mammee apple Clusiaceae 312 254 Mangifera indica L. Mango Anacardiaceae 65; 109 255 Maranta L. Arrowroot Marantaceae 109 arundinacea 256 Marsilea spp. L. Pepperwort Pteridophyta 65 257 Melia azedarach L. Chinaberry tree/ Meliaceae 1 1 3 65; 109; Persian lilac 312; 340 Melia composita Willd. See Melia Meliaceae azedarach 258 Melilotus indica (L.) All. Sourclover Leguminosae- 65; 109 Papilionoideae 259 Melilotus officinalis (L.) Lam. Yellow sweetclover Leguminosae- 1 120 Papilionoideae 260 Mentha citrata Ehrh. Lemon mint Lamiaceae 1 261 Mentha longifolia (L.) Huds. Horse mint Lamiaceae 358 262 Mentha piperita L. Peppermint Lamiaceae 1 2 2 291; 292; 293 263 Mentha spicata L. Spear, wild mint Lamiaceae 1 1 65; 109; 312 264 Mentha spp. L. Mint Lamiaceae 336 265 Mitracarpus scaber Zucc. Rubiaceae 1 2 19; 34; 35 266 Momordica L. Balsam apple/ Cucurbitaceae 34; 35 balsamina southern balsampear Momordica L. See Luffa Cucurbitaceae cylindrica aegyptiaca 267 Momordica spp. L. Momordica Cucurbitaceae 1

81 7 General discussion

Plant species Author Common name Plant family A E O P V Unknown 268 Monarda fistulosa L. Wild bergamot Lamiaceae 1 269 Monodora myristica (Gaertn.) Dunal Calabash nutmeg Annonaceae 1 2 1 270 Mundulea sericea Willd. Silverbush Leguminosae- 65; 109 Papilionoideae 271 Murraya koenigii (L.) Spreng. Currylea f tree/ Rutaceae 1 sweet neem 272 Myristica fragrans Houtt. Nutmeg Myristicaceae 1 273 Myroxylon balsamum (L.) Harms Balsam of Tolu Leguminosae- 1 Papilionoideae 274 Myrrhis odorata (L.) Scop. Anise Apiaceae 291; 292; 293 275 Napoleona imperialis P. Beauv. Lecythidaceae 1 1 276 Nardostachys DC. Indian spikenard Valerianaceae 71 jatamansi 277 Nerium indicum Mill. Scented oleander Apocynaceae 109 278 Nerium oleander L. Oleander Apocynaceae 1 1 95 279 Nicotiana glutinosa L. Tobacco Solanaceae 312 280 Nicotiana rustica L. Rustica, Aztec Solanaceae 284;312 tobacco 281 Nicotiana spp. L. Tobacco Solanaceae 65; 103 282 Nicotiana tabacum L. Cultivated tobacco Solanaceae 3 24; 34; 35; 65; 66; 109; 284; 312 283 Nigella sativa L. Black cumin Ranunculaceae 109;291 284 Ochroma spp. Sw. Ochroma Bombacaceae 65 Ocimum americanum L. See Ocimum Lamiaceae basilicum 285 Ocimum basilicum L. Sweet, forest basil Lamiaceae 1 6 5 65; 66; 68; 103; 109; 114; 180; 291; 292; 293;312 Ocimum canum Sims See Ocimum Lamiaceae basilicum 286 Ocimum gratissimum L. African, shrubby Lamiaceae 3 66; 114 basil 287 Ocimum sanctum L. Holy basil Lamiaceae 1 249 288 Ocimum spp. L. Basil Lamiaceae 34; 35

82 The use ofplan t products to protect stored seeds

Plant species Author Common name Plant family A E O P V Unknown 289 Olea europaea L. Olive Oleaceae 8 290 Opuntia burrageana Britt. & Rose Prickly pear Cactaceae 1 291 Origanum majorana L. Sweet marjoram Lamiaceae 1 Origanum serpyllum Kuntze See Thymus Lamiaceae serpyllum 292 Origanum spp. L. Marjoram Lamiaceae 65; 291 293 Origanum vulgare L. Wild marjoram Lamiaceae 1 2 109 294 Oryza sativa L. Rice Poaceae 1 10 65; 109; 336 295 Pachyrhizus erosus (L.) Urb. Yam bean Leguminosae- 1 312 Papilionoideae Parkia africana R. Br. See Parkia Leguminosae biglobosa Mimosoideae 296 Parkia biglobosa (Jacq.) R. Br. ex G. Nere Leguminosae 1 1 34; 35 Don f Mimosoideae Parkia Keay See Parkia Leguminosae clappertoniana biglobosa Mimosoideae 297 Parkia filicoidea Welw. ex Oliv. African locust bean Leguminosae 26 Mimosoideae 298 Parthenium L. Santa Maria Asteraceae 1 65; 95; 109 hysterophorus feverfew/ carrot grass 299 Pelargonium spp. L'Hér. ex Ait. Pelargonium Geraniaceae 1 300 Peltophorum Sond. Weeping, African Leguminosae- 1 africanum wattle Caesalpinioideae 301 Peltophorum Urb. Leguminosae- 116 suringari Caesalpinioideae 302 Pennisetum (L.) Leeke Pearl millet/ Poaceae 34; 35 americanum American fountaingrass 303 Pennisetum spp. Rich, ex Pers. Kikuyugrass/ Poaceae 2 65; 113; little millet 114 304 Pergularia L. Asclepiadaceae 19; 34; 35; tomentosa 114 305 Petroselinum (Mill.) Nyman ex Parsley Apiaceae 1 291 crispum A.W. Hill 306 Phaseolus spp. L. Bean Leguminosae- 1 65 i Papilionoideae 1

83 7 General discussion

Plant species Author Common name Plant family A E O p V Unknown 307 Phaseolus vulgaris L. Kidney, French Leguminosae- 1 1 109 bean Papilionoideae 308 Physalis minima L. Pygmy Solanaceae 109 groundcherry 309 Phytolacca L'Hér. Pokeweed Phytolaccaceae 167 dodecandra 310 Picrasma spp. Blume bitterwood Simaroubaceae 65 311 Picrolemma sprucei Hook. f. Simaroubaceae 116 312 Piliostigma (DC.) Höchst. Alluba vegetable Leguminosae- 15; 18; 34; reticulatum bush Caesalpinioideae 35; 113; 114; 171 313 Pimenta acris (Sw.) Kostel. Bayrumtree/ Myrtaceae 65; 109 wild clove 314 Pinus spp. L. Pine Pinaceae 1 315 Piper acutifolium Ruiz & Pav. Piperaceae 1 316 Piper guineense Schum. & Thonn. Brown, West Piperaceae 4 5 4 164;336 African pepper 317 Piper nigrum L. Black pepper 1Piperacea e 7 14 1 65; 91; 109; 164; 198; 249; 252; 336 Piper umbellatum L. See Lepianthes Piperaceae peltata \ 318 Pisum spp. L. Pea Leguminosae- 65 Papilionideae 319 Podocarpus nakaii Hay Nakai podocarp Podocarpaceae 109 320 Pogostemon Benth. Patchouli oil Lamiaceae 109 heyeanus 321 Poinsettia (Willd. ex Klotzsch) Poinsettia Euphorbiaceae 109 pulcherrima Graham 322 Polygala butyracea Heckel Polygalaceae 1 323 Polygonum L. Water pepper/ Polygonaceae 1 i hydropiper marshpepper ! knotweed Pongamia glabra Vent. See Pongamia Leguminosae- pinnata Papilionoideae 324 Pongamia pinnata (L.) Pierre Honge/karanjee/ Leguminosae- 1 10 2 65; 109; pongam Papilionoideae 111; 336; 340

84 The use of plant products to protect stored seeds

Plant species Author Common name Plant family A E O P V Unknown 325 Prosopis africana (Guill., Perr. &A . African mesquite Leguminosae 1 153 Rich.) Taub. Mimosoideae 326 Prosopis juliflora (Sw.) DC. Cuji/mesquite Leguminosae 328 Mimosoideae 327 Prunus armeniaca L. Apricot Rosaceae 358 328 Psoralea corylifolia L. Ku tsu/babchi Leguminosae- 1 Papilionoideae 329 Pterocarpus Poir. Barwood Leguminosae- 2 114 erinaceus Papilionoideae 330 Pterocarpus L. f. Red sanders, Leguminosae- 153 santalinus sandalwood Papilionoideae 331 Pulicaria crispa (Forssk.) Benth. ex False fleabean Asteraceae 34; 35 Oliv. 332 Raphanus sativus L. Wild radish Brassicaceae 1 333 Rhazya stricta Decne. Apocynaceae 1 334 Ricinus communis L. Castor Euphorbiaceae 2 17 1 34; 35; 65; 95; 109; 164; 284; 312 335 Rogeria adenophylla J. Gay Pedaliaceae 34; 35 336 Rosmarinus L. Rosemary Lamiaceae 2 2 291; 292; officinalis 293 337 Rottboellia exaltata (L.) L. f. Itch grass Poaceae 19; 34 338 Rumex spp. L. Sorrel/dock Polygonaceae 113 339 Ryania speciosa Vahl Ryania Flacourtiaceae 65; 109; 116; 312 340 Salvadora persica L. Miswak/pilu Salvadoraceae 34; 35; 114 341 Salvia officinalis L. Kitchen sage Lamiaceae 1 1 291; 292; 293 342 Salvia spp. L. Sage Lamiaceae 291 343 Salvia triloba L. Three lobed sage Lamiaceae 291; 292; 293 Sapindus marginatus Willd. See Sapindus Sapindaceae saponaria 344 Sapindus saponaria L. Wingleaf soap berry Sapindaceae 1 65; 109; 358 345 Satureja hortensis L. Summer savory Lamiaceae 1 1 291 346 Saussurea lappa C.B. Clarke Costus Asteraceae 1 65; 109

85 7 General discussion

Plant species Author Common name Plant family A E O P V Unknown 347 Schoenocaulon Sabadilla/ Liliaceae 312 officinale feathershenk 348 Sclerocarya birrea (A. Rich.) Höchst. Jelly plum/ marula Anacardiaceae 1 34; 35 349 Securidaca Fres. Polygalaceae 1 19; 24; 114 longepedunculata 350 Senna occidentalis (L.) Link Septicweed Leguminosae- 1 2 1 24; 34; 35; Caesalpinioideae 168 351 Sesamum indicum L. Sesamum/gingelly Pedaliaceae 23 34; 35; 65; 95; 109; 113; 258; 336 Sesamum orientale L. See Sesamum Pedaliaceae indicum 352 Seseli indicum Wight & Arn. Araliaceae 71; 187 353 Shorea robusta Gaertn. f. Sal tree Dipterocarpaceae 2 336 354 Sida acuta Burm. f. Spinyhead sida/ Malvaceae 1 1 broom weed/ common wire-weed 355 Sida cordifolia L. Country mallow Malvaceae 34; 35 356 Skimmia laureola (Thurb.) Hook. Ner Rutaceae 109 357 Solanum incanum L. Eggplant/nightshade Solanaceae 1 358 Solanum nigrum L. Black night shade Solanaceae 1 359 Sorghum bicolor (L.) Moench Broomcorn Poaceae 1 65 360 Sphaeranthus indicus L. East Indian globe Asteraceae 1 1 thistle 361 Sphenoclea zeylanica Gaertn. Sphenoclea Sphenocleacea 1 362 Spigelia marilandica (L.) L. Woodland pink-root Loganiaceae 109; 116 363 Spirostachys Sond. Tamboti Euphorbiaceae 1 africana 364 Stelechocarpus (Scheff.) R.E.Fr. Annonaceae 1 cauliflorus 365 Stereospermum Cham. Bignoniaceae 15; 18; 34; kunthianum 35; 114; 171 366 Strophanthus A. DC. Arrow poison/ Apocynaceae 1 hispidus hispid strophanthus 367 Strychnos spp. L. Strychnine tree Loganiaceae 284

86 The use of plant products to protect stored seeds

Plant species Author Common name Plant family A E O P V Unknown 368 Swartzia Desv. Pao ferro Leguminosae- 153 madagascariensis Papilionoideae Syzygium (L.) Merr. & Perry See Eugenia Myrtaceae aromaticum aromatica 369 Syzygium cumini (L.) Skeels Java plum Myrtaceae 258 370 Tabebuia rosea (Bertol.) DC. Pink trumpet tree Bignoniaceae 109 371 Tacca (L.) Kuntze Batflower/ Taccaceae 65; 103; leontopetaloides Polynesian 109 arowwood 372 Tagetes minuta L. Stinking roger/ Asteraceae 1 1 65; 167 Mexican marigold 373 Tamarindus indica L. Tamarind Leguminosae- 1 1 65; 109 Caesalpinioideae 374 Tecomaindica Juss. Tecoma Bignoniaceae 109 375 Tephrosia Candida DC. White hoarypea Leguminosae- 109 Papilionoideae 376 Tephrosia spp. Pers. Hoarypea/tephrosia Leguminosae- 65 Papilionoideae 377 Tephrosia vogelii Hook. f. Vogel's tephrosia Leguminosae- 2 95 Papilionoideae 378 Terminalia Guill. & Perr. Almond Combretaceae 34; 35 avicennioides 379 Terminalia sericea Burch. ex DC. Mangwe/almond Combretaceae 1 380 Tetradenia riparia (Höchst.) Codd Lamiaceae 1 381 Tetrapleura (Schum. & Thonn.) Akpa Leguminosae 1 tetraptera Taub. Mimosoideae Thevetia neriifolia Juss. ex Steud. See Thevetia Apocynaceae peruviana 382 Thevetia peruviana (Pers.) K. Schum. Yellow oleander/ Apocynaceae 1 2 23; 95; 116 lucky nut 383 Thujopsis dolabrata Siebold & Zucc. Hiba/mock thuja Cupressaceae 1 384 Thymus serpyllum L. Breckland thyme Lamiaceae 1 1 1 120; 292 385 Thymus vulgaris L. Garden thyme Lamiaceae 1 2 2 120; 291; 292 386 Tiliacordata Mill. Littleleaf linden Tiliaceae 1 387 Tinospora spp. Menispermaceae 139 388 Tragia senegalensis Muell. Arg. Noseburn Euphorbiaceae 23

87 7 General discussion

Plant species Author Common name Plant family A E O P V Unknown 389 Tridax procumbens L. Coatbuttons/ Asteraceae 1 95; 109; Mexican daisy 336 390 Trigonella foenum- L. Sicklefruit Leguminosae- 1 1 65; 109 graecum fenugreek Papilionoideae 391 Verbena officinalis L. Herb of the cross Verbenaceae 1 392 Verbesina (Cav.) Benth. & Golden crownbeard Asteraceae 168 encelioides Hook. f. ex Gray 393 Vigna angularis (Willd.) Ohwi & Adzuki bean Leguminosae- 1 Ohashi Papilionoideae 394 Vigna unguiculata (L.) Walp. Cowpea/ Leguminosae- 1 34; 35 blackeyed pea Papilionoideae 395 Vitellariaparadoxa Gaertn. f. Shea butter tree/ Sapotaceae 3 3 65; 66; karite 103; 109; 114; 164; 171; 336 396 Vitexaltissima L. f. Chaste tree Verbenaceae 1 397 Vitexdoniana Sweet Black plum Verbenaceae 19; 34; 35 398 Vitex negundo L. Begunia/ Verbenaceae 1 2 1 65; 95; negundo chastetree 109; 182; 183; 184; 198; 247; 340 399 Vitexspp . L. Chastetree Verbenaceae 1 400 Warburgia Sprague Pepper-bark Canellaceae 109 ugandensis 401 Weinmannia Cuniniaceae 167 longiflora 402 Withania somnifera (L.) Dunal Winter cherry/ Solanaceae 1 withania 403 Xanthosoma (L.) Schott Cocoyam/arrowleaf Araceae 65; 109 sagittifolium elephant's ear 404 Xeromphis spinosa Thumb. Keay Kucuruman Rubiaceae 109 405 Ximenia americana L. Tallow wood Olacaceae 15; 18; 114; 171 406 Xylocarpus (Imkhan.) Roem. Cannonball tree Meliaceae 109 moluccensis 407 Xylopia aethiopica (Dunal) A. Rieh. West African Annonaceae 2 1 pepper tree/ Ethiopian pepper

88 The use of plant products to protect stored seeds

Plant species Author Common name Plant family A E O P V Unknown 408 Zanthoxylum alatum Roxb. Wingleaf prickly Rutaceae 1 109 ash 409 Zanthoxylum (Lam.) Zepern. & Rutaceae 1 3 zanthoxyloides Timler 410 Zea mays L. Maize/corn Poaceae 12 34; 35; 65; 109; 312; 336 411 Zingiber cassumunar Roxb. Cassumunar ginger/ Zingiberaceae 62 plai 412 Zingiber officinale Roscoe Garden ginger Zingiberaceae 4 164; 258; 340 413 Zingiber spectabile Griff. Black gingerwort Zingiberaceae 1 414 Zingiber zerumbet (L.) Sm. Bitter ginger Zingiberaceae 1 415 Cattle dung 2 26; 65; 109; 194 416 Daddoya Acanthaceae 270 417 Dalda 1 418 Goga masu 270 419 Wood 11 15; 16; 18; 19; 23; 26; 34; 65; 91; 109; 113; 114; 119; 151; 153; 208; 258; 270; 312; 339; 354

*: A = ashes, E = extracts, O = oils (non-volatile), P= powder or fresh, U= unknown #, V = volatile oil, numbers indicate the number of references mentioned in the separate tables. #: Application, quantity, beetle species and/or effect are not noted in the references.

89 7 General discussion

90 Theus eo fplan tproduct st oprotec tstore dseed s

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1 Abdul Kareem A., 1980.Nee m as an antifeedant for certain phytophagous insects and abruchi d on pulses.Proceeding s of the First International Neem Conference, Rottach-Egern, pp 223-250. 2 Adebayo T.A. &Gbolad e A.A., 1994.Protectio n of stored cowpea from Callosobruchus maculatus using plant products. Insect Science and its Application, 15:185-189 . 3 Adedire, CO., Lajide L., 1999.Toxicit y and oviposition deterrency of some plant extracts on cowpea storage bruchid, Callosobruchusmaculatus Fabricius. Zeitschrift für Pflanzenkrankheiten und Pflanzenschutz, 106: 647-653. 4 Ahmed S.M. &Ahama d A., 1992.Efficac y of some indigenous plants as pulse protectants against Callosobruchuschinensis (L.)infestation . International Pest Control, 34:54-56 . 5 Ahmed, K.S.,Haqu e M.A. &Isla mB.N. , 1993.Efficac y of edible and non-edible oils against the pulse beetle, Callosobruchus chinensis (Coleoptera: Bruchidae). Bangladesh Journal of Entomology, 3: 1-7. 6 Ahmed K., Khalique F.,Afzal M., Malik B.A. &Mali k M.R., 1988.Efficac y of vegetable oils for protection of greengram from attack of bruchid beetle. Pakistan Journal of Agricultural Research, 9:413-416. 7 Ahn Y.J., Lee S.B., Lee H.S.& Ki m G.H., 1998.Insecticida l and acaricidal activity of carvacrol and ß-thujaplicine derived from Thujopsisdolabrata var. hondai sawdust. Journal of Chemical Ecology, 24: 81-90. 8 Ajayi O., Arokoyo J.T., Nezan J.T., Olayinka O.O.,Ndirmbul a B.M. &Kannik e O.A., 1987. Laboratory assessment of the efficacy of some localplan t materials for the control of storage insectpests . Samara Journal of Agricultural Research, 5: 81-86. 9 Ajibola Taylor T., 1975.Effect s of orange and grapefruit peels on Callosobruchusmaculatus infestation of cowpea.Ghan a Journal of agricultural Science, 8: 169-172. 10 Al-Hemyari A.A, 1994. Effectiveness of someplan t products as faba-bean protectants against cowpea beetle, Callosobruchusmaculatus F. (Coleoptera: Bruchidae). Annals of Agricultural Science, Moshtohor, 32:997-1007 . 11 Ali S.I., Singh O.P. &Misr a U.S., 1983.Effectivenes s ofplan t oils against pulse beetle Callosobruchus chinensis Linn. Indian Journal of Entomology, 45: 6-9. 12 Allotey J. &Dankwa h J.A., 1995.Som e aspects ofth ebiolog y and control of the cowpea weevil, Callosobruchus maculatus (F.) onbambar a groundnut and cowpea. Insect Science and its Application, 16:223-228 . 13 Alzouma I., 1989.Economi c incidence of sahelian cowpea bruchids in Africa. Propositions for controlling this grain legume pest. In: Yoshida T. (Ed.). Proceedings of the Second International Symposium on Bruchids and Legumes, Okayama, Japan, pp 1-14. 14 Alzouma I., 1995.Connaissanc e etcontrôl e des coléoptères Bruchidae ravageurs des légumineuses alimentaires au Sahel. Sahel IPM, 1:2-16 . 15 Amadou B., 1985.Effet s de quelques plantes insecticides et (ou)insectifuge s sur la réproduction et le développement deBruchidius atrolineatus Pic. et Callosobruchus maculatus (Coleoptera Bruchidae) ravageurs du niébé (Vigna unguiculata(L. ) Walp). Diplôme d'Agronomie Tropicale, Filière:Protectio n des Cultures,Ecol e Supérieure d'Agronomie Tropicale deMontpellier , 54pp . 16 Amuti K. &Larb i M., 1981.Postharves t losses inth e bambara and geocarpa groundnut seeds stored under traditional conditions in Ghana.Tropica l Grain Legume Bulletin, 23:20-22 . 17 Anonymous, 1976.Contro l ofeowpe a weevil in storage.Tropica l Grain LegumeBulletin , 6:36 . 18 Anonymous, 1984.Le sbruche s du niébé -dégât s -moyen s de lutte;méthode s traditionnelles de conservation en grenier. Fiche Technique Vulgarisation, INRAN Département Recherches Agricoles Section Protection desVégétau x Entomologie, Niamey, Niger, 4pp . 19 Anonymous, 1991. Enquête sur le stockage du niébé en milieu villageois auNiger . Département de Biologie de l'Université deNiamey , Département de Formation en Production desVégétau x du C.I.L.S.S., 8pp . 20 Anonymous, 1991. Les ennemis des cultures vivrières dans le Sahel.C.I.L.S.S. , CTA ,p p 6-9. 21 Anonymous, 1995. Conservation duniéb é avec l'huile deneem . Fiche Technique SPV GTZ, Benin, 21pp .

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44 Caswell G.H., 1981. Damage to stored cowpea in thenorther n part ofNigeria . Samaru Journal of Agricultural Research, 1: 11-19. 45 Chander H.& Ahme d S.M., 1982.Extractive s ofmedicina l plants aspuls eprotectant s against Callosobruchuschinensis L. infestation. Journal of Food Science and Technology, 19:50-52 . 46 Charjan S.K.U. &Tarar , J.L., 1994.Viability , vigour andmycoflor a changes in pulse beetles damaged seeds of greengram (Vignaradiata L .Wilczek) .Proceeding s ofth eNationa l Academy of Sciences, India. Section B,Biologica l Sciences, 64:95-98 . 47 Chinwada P. &Gig a D., 1993.Vegetabl e andnee m oils asprotectant s of stored beans against bruchids. Second meeting of the Pan-Africa working group on bean entomology, Harare, Zimbabwe. CIAT-African-Workshop-Series, 25:40-49 . 48 Chinwada P.& Gig a D.P., 1996.Sunnin g asa technique for disinfesting storedbeans . Postharvest Biology and Technology, 9: 335-342. 49 Chinwada P. &Gig a D.P., 1997.Traditiona l seedprotectant s for the control ofbea n bruchids. Tropical Science, 37:80-84 . 50 Chiranjeevi C. & Sudhakar T.R., 1996.Effec t of indigenousplan t materials onth e fecundity, adult emergence and development ofpuls ebeetle , Callosobruchuschinensis (L.)i nblackgram . Journal ofResearc hA PAU , 24: 57-61. 51 Chiu S.F., 1939.Toxicit y studies of so-called "inert"material s with the bean weevil, Acanthoscelides obtectus (Say).Journa l of Economic Entomology, 32:240-248 . 52 Cockfield S.D., 1992.Groundnu t oil application and varietal resistance for control of Callosobruchus maculatus(F. ) in cowpea grain inTh e Gambia. Tropical Pest Management, 38: 268-270. 53 Cortesero A.M., Monge J.P. &Huignar d J., 1997.Dispersa l andparasitizin g abilities oiEupelmus vuilleti (Hymenoptera: Eupelmidae) within a column of cowpea seeds. Environmental Entomology, 26: 1025-1030. 54 Couilloud R., 1991. Insectes, araignées &acariens .Correspondance s entre les dénominations scientifiques et anglo-saxonnes. Centre de Coopération Internationale en Recherche Agronomique pour le Développement, 681pp . 55 Credland P.F., 1992.Th e structure ofbruchi d eggsma y explainth e ovicidal effect ofoils . Journal of Stored Products Research, 28: 1-9. 56 Credland P.F., Dick K.M. &Wrigh t A.W., 1986.Relationship s between larval density, adult size and egg production in the cowpea seedbeetle , Callosobruchusmaculatus. Ecological Entomology, 11: 41-50. 57 Cruz C.& Cardona E., 1981. Control of dry seed weevilswit h cooking oil.Journa l of Agriculture of University of Puerto Rico, 65:295-298 . 58 Dabire L.B., 1985.Le s structures et lestechnique s traditionnelles de conservation duniéb é au Burkina Faso.Note s et documents Burkinabés: Bulletin Trimestriel d'Information Scientifique et Technique, Ouagadougou, Burkina Faso, 16:51-63 . 59 Dabire L.B., 1985.Le s méthodes traditionnelles deprotectio n duniéb é contre lesbruche s au Burkina Faso.Note s et documents Burkinabés: Bulletin Trimestriel d'Information Scientifique et Technique, Ouagadougou, Burkina Faso, 16:64-77 . 60 Dabire C, 1991. Lesméthode straditionnelle s deprotectio n des stocks de niébé(Vigna unguiculata) auBurkin a Faso.Atelie r Régional Ouest-Africain sur laProtectio n Naturelle des Végétaux, avec l'Appui d'AGRECOL, Madesahel, Mbour, Sénégal.End a t.m, 6pp . 61 Dabire C, 1992.Le s méthodes traditionnelles deprotectio n des stocks deniéb é auBurkin a Faso. SahelPV Info, 49:7-13 . 62 Dadang, Riyanto S. &Ohsaw a O., 1998.Letha l and antifeedant substance from rhizome oîAlpinia galanga Sw. (Zingiberaceae). Journal of Pesticide Science, 23:304-307 . 63 Daniel S.H. & Smith R.H., 1991.Th e repellent effect ofneem (Azadirachta indicaA .Juss. ) oil and itsresidua l efficacy against Callosobruchusmaculatus (Coleoptera: Bruchidae) on cowpea. In: Fleurat-Lessard F. &Duco mP . (Eds.).Proceeding s ofth e 5th International Working Conference on Stored Product Protection, Bordeaux, France,p p 1589-1597. 64 Das G.P. &Kari m M.A., 1986.Effectivenes s ofneem seed kernel oil as surface protectant against the pulse beetle, Callosobruchuschinensis Linn. (Bruchidae: Coleoptera). Tropical Grain Legume Bulletin, 33:30-33 .

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