BASE Biotechnol. Agron. Soc. Environ.201216(2),269-276 Focus on:

SilkmothsinMadagascar:Areviewofthebiology,uses,andchallengesrelatedtoBoroceracajani(Vinson,1863)(Lepidoptera:Lasiocampidae)TsiresyM.Razafimanantsoa(1),GabrielleRajoelison(2),BrunoRamamonjisoa(2),NoromalalaRaminosoa(1),MarcPoncelet(3),JanBogaert(4),ÉricHaubruge(5),FrançoisJ.Verheggen(5)(1)UniversitédeAntananarivo.FacultédesSciences.DépartementdeBiologieanimale,ÉcologieetConservation.BP906.Antananarivo(Madagascar).(2)UniversitédeAntananarivo.ÉcolesupérieuredesSciencesagronomiques.DépartementdesEauxetForêts.BP175.Antananarivo101(Madagascar).(3)Univ.Liège.DépartementdeSociologieduDéveloppement.BoulevardduRectorat,7.B-4000Liège1(Belgique).(4)UniversitélibredeBruxelles.Serviced’ÉcologieduPaysageetSystèmesdeProductionvégétale.AvenueFranklinRoosevelt,50.B-1050Bruxelles(Belgique).(5)Univ.Liege-GemblouxAgro-BioTech.Unitéd’Entomologiefonctionnelleetévolutive.PassagedesDéportés,2.B-5030Gembloux(Belgique).E-mail:fverheggen@ulg.ac.be

ReceivedonDecember3,2010;acceptedonDecember8,2011.

Borocera cajani or “Landibe” (vernacular name) is the wild silk moth that is currently used to produce silk textiles inMadagascar.ThisspeciesisendemictoMadagascar,andisdistributedthroughouttheisland,colonizingtheUapaca bojerior“Tapia”forestofthecentralhighlands.TheforestprovidesfoodintheformofplantsforB. cajani,includingU. bojerileaves.Thespeciessecretessilkattheonsetofpupationandformakingcocoons.Borocera cajanianditsnaturalhabitatarethreatenedbyhumandestruction,suchasbushfires,firewoodcollection,charcoalproduction,andtheover-harvestingoftheircocoons.Wildsilkproductionlargelydisappearedwhenthesilkindustryutilizedmanypeopleontheislandasthecollectorsofcocoons,spinners,dyers,weavers,andartistswhotransformthesilkintoclothes,accessories,andobjects.Therefore,itisimportanttostudythebiologyofB. cajanitorevitalizesilkproductioninawaythathelpsconservethisspeciesandtheTapiaforest.Keywords.Borocera cajani,Uapaca bojeri,silk,silkindustry,biodiversity,Madagascar.

Les vers à soie endémiques de Madagascar : synthèse bibliographique de la biologie, des utilisations et des perspectives associées à Borocera cajani (Vinson, 1863) (Lepidoptera : Lasiocampidae). Borocera cajani ou «Landibe» (nomvernaculaire)estl’undespapillonsséricigènessauvagesdontlasoieestlaplusutiliséedansledomainetextiledeMadagascar.Cette espèce endémique s’observe dans toute l’île,mais colonise particulièrement la forêt deUapaca bojeri ou forêt de«Tapia» des hautes terres centrales. La forêt fournit les aliments àB. cajani tels que les feuilles deU. bojeri. L’espècesecrètelasoiequandelleentreennymphoseetconstruitsoncocon.Borocera cajanietsonhabitatnaturelsontmenacésparlesdestructionsde l’Hommetellesque les feuxdebrousse, lacollectedeboisdechauffage, laproductiondecharbondeboisetlasurexploitationdeleurscocons.Laproductiondesoiesauvageestenconstantediminution,alorsquelafilièresoieimpliquebeaucoupdegensdansl’îlecommelescollecteursdescocons,lesfileurs,lesteinturiers,lestisseursetlesartistesquitransformentlasoieenhabits,enaccessoiresetobjets.Revitaliserlafilièresoieestunmoyendefavoriserlaconservationdecetteressourcenaturelleetdesonhabitatforestier.Mots-clés.Borocera cajani,Uapaca bojeri,soie,industriedelasoie,biodiversité,Madagascar.

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1. INTRODUCTION

Madagascarisoneofthemostimportantcentersofworldbiodiversity, with a high level of endemism, around90% (Mittermeier et al., 2004).Many of the speciesare forest dwelling (Myers, 1988), with biodiversityencompassingallclassesofanimals,includinginsects.Among these insects, moths are used in the textileindustry.Madagascan textile silk has a long history.Indeed, wild silk, termed “Landibe”, was exploitedlong before the introduction of “Chinese silk” fromBombyx moriduringthe19thcentury(Rafidiarimalala,1974;Costa,2004).WildsilkinMadagascarhasmanyorigins,withmanysilkproducingspeciesbeingpresenton the island. One such species is Borocera cajani(ONU,1991).Historically,theMadagascanpopulationused B. cajani silk to create sumptuous shrouds.However,thecontinueduseofthissilk,alongwiththegradualdisappearanceofnativeforests,hasnegativelyaffected thedistributionandpopulation levelsof thisspecies. Therefore, it is necessary for researchers tostudy thebiologyofB. cajani to safeguard its future(Paulian,1953).

2. BIOLOGY

2.1. Taxonomy

The genus Borocera was originally described byBoisduval(1833),andthespeciescajaniwasdescribedby Vinson (1863). This species has been confusedwithBorocera madagascariensis,whichoftenresidesalongthecoastalpartoftheisland.TheB. cajanisilkmothbelongstotheLasiocampidaefamily,andtotheGonometinae subfamily. This subfamily is endemictoMadagascar,andincludesthelargerspeciesof theLasiocampidae(DeLajonquière,1972;ONU,1991).Itischaracterizedbyveineightofthehindwings,whichconnectstoveinsevenatadistancefromitsbase,andthen forms a secondary cell of length that is nearlyequal to the top of the median cell, with both cellsbeingaboutthesamewidth(Aurivillius,1927).

2.2. Morphology

The genus Borocera encompasses all moths withfasciculate antennae,which areopenlyuneven alongthefirstthirdofthesegmentfromthebase.Mothsofthisgenusalsohavea smallhead,noproboscis, andsmall,slightlyprominent,eyes.Borocera cajaniisthemost widespread moth in Madagascar, and withoutdoubtisoneofthemostabundant.Ithasmanyformsandlandraces(DeLajonquière,1972).

Both sexes exhibit very high dimorphism in sizeand color. The genitalia remain however constant,

regardless of the origin of species being examined(De Lajonquière,1972).Thewingsaregenerallydarkand fuzzy for the male and a uniform grey for thefemale (Paulian, 1950; Razafindraleva, 2001).Adultmales and females have several main characteristics(Figure 1).

Malesaresmallish,hairy,andthick-bodiedmoths.Their antennae are well developed and bipectinate.Their head, thorax, and legs are the same color asthe forewing. The wingspan is about 38-52mm(De Lajonquière, 1972). The forewings are oblongandwelldevelopedinwidth,whiletheapexisslightlysinuous or round. Their color is red ochracea tobrown,withasinuatesubmarginal line,apostmedialtransverse line, and a stigma grayish point in thediscalcell(Paulian,1951).Theformof theforewingvaries for one egg-laying (Paulian, 1953).The colorof the hindwings is quite similar to the forewings,but is often overshadowed in part or inwhole. Liketheforewings,thedorsalwingsaresometimesbarredtransverselywithanapparentmedianreddishshadow,and are sometimes obsolete (De Lajonquière, 1972;Razafindraleva,2001).Thegenitaliaare formedbyamembrane that leads to a thicker penis and serratededges.Thepenisiscrossedbyaprominentejaculatoryduct.Twovalvesarelocatedoneithersideofthepenis(DeLajonquière,1972;Razafindraleva,2001).

Females aremuch larger;with a body size aboutthreetimesthatofthemale.Theirwingspanisabout70-75mm, with a forewing length of 35mm. Theantennalshaftisblack,withyellowpectination.Thesemoths are hairy,with a thick bodied, grayish yellowtinted,bare thorax,and legsof thesamecoloras theforewings.Thehairylegsterminateinblacktarsi.Thewingsareadirtywhite topalegrey,silky,andshiny(De Lajonquière, 1972; Razafindraleva, 2001). Theforewingsarecrossedbytwotransverselinesthatspilt

Figure 1. Borocera cajani moths (left: female, right:male)—Papillon de Boroceracajani (à gauche : femelle, à droite : mâle).

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thewingintodifferentspaces(Razafindraleva,2001).Thediscalareahasadarklunuledotattheendofthecell. The color of the hindwings is the same as theforewings.Thebackofthefourwingsisyellowishsepiaincolor,whichisconsistentwithmuchlessapparentmedian gray shadows. The apex of the abdomen ischaracterized by two well developed thick ridges,withsensoryhairs.Thegenitalopeningisformedbythesuperpositionoftwobeads.Thisstructureensurescoaptationduringmatingandovoposition.

3. LIFE CYCLE AND HABITAT

Borocera cajani isbivoltine,meaning that ithas twogenerations inayear (Paulian,1953;Razafindraleva,2001;Razafimanantsoa,2008).Thefirstgenerationofeggshatchesduringtherainyseason,betweenJanuary(Razafimanantsoa,2008)andMarch (Razafindraleva,2001).ThesecondgenerationofeggshatchesbetweentheendofAprilandNovember(Razafindraleva,2001;Razafimanantsoa, 2008). The life cycle of the silkmoth comprises four stages: the egg, caterpillar orlarva, chrysalis or pupa, and adult.After emergence,femalesattractmalesbyreleasingasexualpheromone(Zborowski et al., 2007). For both sexes, emergenceprimarilyoccursbetweentwoandsixintheafternoon.Copulationislargelyobservedbetweenoneandsevenintheafternoon,lastinguptoonehour.Onemalemaymatewithuptothreefemales.Theadultdoesnotfeed.Eggsarelaidfromsevenintheafternoononwardsofthesamedaythattheadultfemaleemergedandmated.Unfertilefemalesmaylayeggs,buttheseeggsdonothatch.The number of eggs that are laid varies from250to845.Alleggsarelaidontheleaves,branches,and trunk surface of specific host plants (includingU. bojeri) (Razafimanantsoa, 2008). Females layeggs in captivity, even without the necessary hostplants.TheeggsofB. cajaniareabout2mminsize,quite hemispherical, with a hard shell and granulartexture.Theporcelain-likeeggsarecoveredinagrey-greenishband.Themostfertileeggshatchafterabout10days,withahatchingsuccessrateofabout89%.NoparthenogeneticcharacteristicshavebeenobservedforB. cajani.

Thehighestmortalityoccursduringthefirstinstar(Paulian, 1953).At this stage, the larvaedisperse bydropping from twigs on a long silken thread (Piney,1975),butwhichmayresultintheirlandingonaplantofthewrongfoodtype.Ifthefirstinstardoesnoteatwithin4daysofhatching, itcannotsurvive(Paulian,1953).Asthecaterpillarsdevelopthroughfivestages,theychangeinsizeandshape.Thesecondinstarlarvaedevelopfourspinybouquetstowardthethoracicregion.Thespinybouquetsprogressivelydevelopinsizewitheachinstarand,whenalarmed,thecaterpillarsexpose

four“pom-pom”likespinybouquets,whichareredatthebaseandblackontop.Somelarvaeshowonlytheblackishbristle.Thefifthinstarcaterpillar(Figure 2)isveryhairyatthelateralface,andisimmense,reachingover 12cm long (Grangeon, 1906;Razafimanantsoa,2008;Rakotoniaina,2009).Inthewild,thefifthinstarisremarkablywellcamouflagedtomimicthebarkoftheirplantfood.Theircolorisgrey-blue.

ThecocoonofB. cajaniisovalinshapeandgrayishincolor,andprojectsurticant(i.e.,itchy)hairs.Thesizeoffemalecocoonsisabout50mminlength,andabout25mmindiameter,andissmallerformales.Thecocoonhasthreelayers,weighing300-400mgforfemalesand90-200mg for males. The prepupa stage lasts 3to 5days(Grangeon,1906;Razafimanantsoa,2008).

As with many moths, B. cajani is preyed on bymanypredatorsandissubjecttomanydiseases,withall stages being at risk of attack (Zborowski etal.,2007). Bird predators include Acridithores tristis,Cuculus rochii, Centropus toulou, Corvus alba,and Hypsipetes madagascariensis. However, otherpredatorsalsoexist, includingbats, reptiles (Furcifer lateralis), the praying mantis, ants, and spiders.Parasitesare themost invasive,andmostlycomprisewasps (Ichneumonidae:Pimpla,Xanthopimplahova,Brachymeria borocera, Ophion sp., Braconidae:Apanteles boroceraeandApanteles decaryi).Anotherinvasive species is Synthesiomyia nudiseta (Diptera:Tachinidae), which, while it attacks the larvae, maystill emerge in a subsequent stage (Grangeon, 1906;Grangeon,1907;Paulian,1953;Razafindraleva,2001;Razafimanantsoa,2008).Thereisalsoawiderangeoffungal,bacterial,andviraldiseases,suchas“pebrine,muscardineandflasherie”(Grangeon,1907).

ThelifecycleofB. cajanirequires102to193daystocomplete (Razafindraleva,2001;Razafimanantsoa,2008) (Figure 3). This major variation in the daysrequired to reach the pupal stage is due to diapauseduringthedryseason.

Borocera cajani colonizes the U.bojeri (or“Tapia”)forest,whichisfoundinthecentralhighland

Figure 2.Borocera cajani larvae in5th instar—Larve de stade 5 deBoroceracajani.

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zones(Imamo,Itremo)andIsalozonesofsouthwestof Madagascar (Gade, 1985). The “Tapia” forestconstitutesoneofthenaturallyformedforestsoftheisland, primarily containing just one tree species,U. bojeri (Rakotoarivelo, 1993;figure 4).This plant

isoneofthemajorplantfoodsofB. cajani.Thesilkmothsareverypolyphagous,andarefoundinhabitingmanyhabitatsofMadagascar,includingDiegoSuerez,Montagned’Ambre,NosyBe,Anjozorobe,Analavory,Ankazobe region, Antananarivo, Manjakatompo,Ankarana’s massif, Ambohimahasoa, Andringitra’smassif, Bekily, Majunga, Ankarafantsika’s Region,Antsingy, Antsalova region, and Morondava(De Lajonquière, 1972). They also feed onPsidium guajava L., Psidium cattleianum Sabine, Aphloiatheiformis(Vahl)Benn.,Dodonaea madagascariensisRadlk., Cajanus indicus Spreng., Harunganamadagascariensis Lam. ex Poir., Acacia dealbataLink,Schefflerasp.J.R.Forst.&G.Forst.,Eucalyptussp.L’Hér.,Tamarindusindica L.,Avicennia officinaliL., Salíx babylonica L., and Terminalia catappa L.(Paulian,1953;DeLajonquière,1972;Rakotoarivelo,1993;Razafindraleva,2001;Razafimanantsoa,2008;Rakotoniaina,2009).

The herbaceae stratum in the “Tapia” forest hasan important role in the life cycleofB. cajani.Thisstratumservesasplaceforchrysalisnidification(i.e.,chrysalisconstruction)(Razafimanantsoa,2008).

Figure 3.Borocera cajanilifecycleininsectariuminAmbohimanjakain2006—Cycle de vie deBoroceracajaniélevé en insectarium à Ambohimanjaka en 2006.

Figure 4. Uapaca bojeri forest in Arivonimamo,2009—Forêt de UapacabojeriàArivonimamo,2009.

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4. IMPORTANCE

TheGonometinaespeciesarecurrentlyusedforwildsilk production in Madagascar, with the silk beingprimarilyspunfromB. cajani(DeLajonquière,1972).The size of the harvest varies greatly from year toyear,reaching10tto43tforthewholeisland(Gade,1985;CITE/BOSSCORPORATION,2009).Wildsilkproductionisasourceofincomeforthecommunitiesliving adjacent to theTapia forest (Kull, 2003). Silkproductionprovidesanalternativesourceofincometothepoorestandmostdisadvantagedpeople,especiallyduring“leanperiods”(Kulletal.,2005).Thevillagersgathercocoonsandsellthem.Thechrysalisisremovedandeaten,whiletheemptycocoonsarecooked,spun,andwoven into silk fabrics (Kull,2003).Thirty-fourpercentofhouseholdsinAmbohimanjakaand65%ofhouseholdsinIlakaearncashincomesfromtheharvestof B. cajani cocoons (Kull, 1998; Kull, 2002). Thelargerrainyseasonsilkharvestprovidesanimportantcash income during the meager months before thericeharvest.CollectingBoroceracocoons,processingthem,unravelingthefilamentstospinintothread,andweavingthemintosilkhavebeenpart-timeactivitiesofhighlandricegrowingpeasantsforseveralhundredyears(Gade,1985).

SilkproductioninvolvesmanyMadagascanpeople,from the wild silk harvesters to the spinners, dyers,weavers, and craftsmen who transform the silk intoclothing,accessories,anddecorativeitems.

Borocera pupae are also a preferred delicacy.Besidesanimalprotein,theinsectsprovideoccasionalvariation in textureand taste to thedailydiet,whichprimarily constitutes rice andmanioc (Decary, 1937;Gade,1985;Guigou,1989;Kulletal.,2005).

Borocera cajanisilkisprocessedtoproduceritualburial shrouds and clothing items throughout thehighlandsaspartofthearea’sculture.The“Landibe”shroudisessentialforanyrespecteddeadperson.Tobewrappedbymany“Lambamena”(redsilkshrouds)isasignofsupremehonoraccordingtotheMadagascanadage:“Izay sahy maty mifono lambamena”(literallytranslated as “the intrepid facing the death wearsone “Landibe” shroud”) (Rakotoniaina, 2009). The“Landibe” shroud is preferred over that made withBombyx mori for Madagascan ritual burials andfor exhumations or “Famadihana” (a Madagascanhabit to renew the shroudofmortal remains every5to 10years) (CITE/BOSS CORPORATION, 2009)(Figure 5).Indeed,wildsilkispreciousforthissecularusage (Rakotoniaina, 2009), even if few people canaffordthistypeofcloth.Theclothcosts25to71Euros(CITE/BOSS CORPORATION, 2009). The silkproduced fromB. cajani cocoons is very remarkableforitsglow,tenacity,elasticity,andsolidity(Coquerel,1854;Guigou,1989).Forthisreason,theMadagascan

peopleseekthissilkoverothersilks.However,somepeoplealsouseBombyx moriforshrouds.

5. PROBLEMS

Estimatesshowthatbetween150,000and200,000haofMadagascanforestislostperyear(Mintenetal.,2003).Povertyisonerootcauseofthelossofbiodiversity.AswithmanyspeciesinMadagascar,B. cajaniiscriticallyendangeredduetothedestructionoftheirnativeforest(Razafimanantosoaetal.,2006). In1933, the Imamo“Tapia”forestwasviewedasbeingimportant,andwasrecognized for the collection of cocoons. However,deforestation over the subsequent 20years (until the1950s) resulted in thisareabecominga sparse forest(Paulian, 1953). Consequently, since the 1960s, thiszone ceased to produce “Landibe” (Razafintsalamaetal.,1999;Kull,2003;Kulletal.,2005).TheIsaloand Itremo zones remain the producing centers ofwild silk in Madagascar (Paulian, 1953). However,Madagascanwildsilkproductioncontinuestodecline.In1902,theIslandproduced103tofemptycocoonsversusanestimated43twasproducedin2009(CITE/BOSSCORPORATION,2009).Inaddition,inItremo,cocoonsareonlyabundantincyclesofoneyearoutoffive.Asaresult,thesparsecocoonpopulationsarenotworththeharvesteffort(Gade,1985).Therearemanycontributorstothedeclineofthesilkmoth,andhencewildsilkproduction,including:– thedestructionofpreferredmothhabitatscausedby thewaythatharvestersgathercocoonsintheforest bybreakingmanybranches(Grangeon,1906);– the proliferation of introduced species, such as Pinus sp., which outcompetes native plants, such asU. bojeri, by its quick growth and alteration of microhabitats due to its heliophile characteristics, leadingtochangesinsoilquality(Kulletal.,2005);– bushfires,whichkillB. cajani anddestroyyoung U. bojeri trees, seedlings, and sprouts (Grangeon, 1910; Perrier de la Bâthie, 1921; Paulian, 1953; Vignal,1963;Gade,1985;Guigou,1989;Kulletal., 2005);– theoverharvestingofpupae for food consumption (Paulian,1953;Kulletal.,2005;Razafimanantosoa etal.,2006;Razafimanantsoa,2008).

Thedemiseoftheindustryhasalsobeenaffectedbythelackofmodernizationofnativesilkmanufactureandthehighcostof“Landibe”burialshrouds.About53tperyearofsilkisproducedfromthedomesticatedChinese silkworm Bombyx mori (CITE/BOSSCORPORATION,2009).However,thepriceofthreadandclothesmadewiththissilkisalmostthesameasforB. cajaniproducts.Bombyx moriisproducedwithcheap labor, as this species can be easily reared in

274 Biotechnol. Agron. Soc. Environ. 201216(2),269-276

largequantities.Thisresultsinanestheticallysuperiorfabric,whichisexportedinamarketwhereB. cajanidoesnotcompete(Rafidiarimalala,1974;Gade,1985).The “Landibe” cocoons cannot be spun. It must beskinned off its chrysalis by cocoon incision. Thespinning-millismadeusingdistaff,andproductivityislimited(Rakotoniaina,2009).

ManagementmethodsincludingGELOSE(“Gestion locale Sécurisée,”meaningsecurelocalmanagement)and GCF (“Gestion Contractualisée des Forêts,”meaningcontractualmanagementofforests)arenowentrustedtolocalbasedcommunitiestomanagelargepartsofthe“Tapia”forest.Localbasedcommunities,withlegalconstitutions,arecomposedofresidentsinahamlet,village,orgroupofvillages.Thesevolunteersare united by common interests and are willing toobey the rules of common life, and run the naturalresource management associations (GELOSE, 1999;Robsomanitrandrasana,2008).Theirrole is toensure

the self-sustainablemanagement of natural resourcesthatarepresentontheirterritory,includingtheforestand fauna, such as the silkworm (Razafindrakoto,2005;Robsomanitrandrasana,2008).Currently,duetocommunitymanagement,conservation,andrecoveryoftheforest,theseorganizationshaveprovedasuccessfultransfer of good management, with the cessation ofcharcoal production (Consortium RESOLVE/PCP/IRD, 2005; Robsomanitrandrasana, 2008). However,somevillagersdonotwishtojointheassociation(localcommunitybased)toavoidfollowingtherules.

6. CONCLUSION AND RECOMMENDATIONS

AlthoughsericulturecontinuestoplayaminorroleinMadagascar at an international level (Krishnaswamiet al., 1974), it remains a limited source of incomefor local people who depend on it. More than

Figure 5.Wrappingduringthe“Famadihana”inArivonimamo,2009—Enveloppement de dépouilles mortelles pendant le « Famadihana » àArivonimamo, 2009.a:mortalremainsoutofthegrave—dépouilles mortelles exhumées;b:shroudmadebyBombyxmori—linceul fait à partir de Bombyxmori;c:wrapping—enveloppement;d:dancewithmortalremainswrappedinsilkshrouds—danse avec les dépouilles mortelles placées dans les linceuls en soie.

a

c

b

d

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10,000familiesworkinthesilkindustry(CITE/BOSSCORPORATION,2009).Therefore,itsrevitalization,alongwithcarefulmanagement,couldbeanimportantfactorforthefutureconservationofsilkmoths,suchasB. cajani.Inrecentyears,humanpressuresonB. cajaniand its habitat have decreased through applyingmanagement methods led by local communities.However, it remains a challenge to convince peoplewhodonotwishtojointheassociations,tomaintainthesamelevelofmanagementandthusconservation,toensureeveryonehasthesameconviction.Therefore,acomprehensivestudyonthebiologyandecologyoftheinsectanditshabitatisnecessarytoestablishlong-termmanagementprotocols.

Thepermanentcessationofbushfirepracticeandlogging,thereforestationofU. bojeri,andeliminationofintroducedplants(suchasPinus)withinthe“Tapia”forestareveryimportantactionstowardsmaintainingthe natural habitats of B. cajani (ONU, 1991;Razafimanantsoa,2008).Restrictionson themethodsof collecting cocoons, such as avoiding breakingbranches, and the timing of cocoon gathering, toensuretheestablishmentofthenextgeneration,mustbeadheredto.TherepopulationofB. cajaniinthewildrequires an understanding of the various biologicaland human impacts on the population for successfulproduction. The provision of information throughenvironmentaleducationtothechildrenoftheregionwhereB. cajaniisabundantwouldhelpsafeguardthefuture of this industry.Through thismechanism, theMadagascanpeoplemaydevelopanunderstandingoftheimportanceofB. cajani asaculturalheritage.

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