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ExperimentalandAppliedAcarology23:659667,1999.

1999KluwerAcademicPublishers.PrintedintheNetherlands.

AcomparativestudyofVarroajacobsonireproductionin
workercellsofhoneybees(Apismellifera)inEngland
andAfricanizedbeesinYucatan,Mexico
LUISMEDINAMEDINA1andSTEPHENJOHNMARTIN2*
1

DepartamentodeApicultura,FacultaddeMedicinaVeterinariayZootecnia,UniversidadAutonoma
deYucatan,ApartadoPostal4116,C.P.97100,Merida,Yucatan,Mexico; 2NationalBeeUnit,Central
ScienceLaboratory,SandHutton,York,YO411LZ,UK
(Received10March1999;accepted2June1999)

Abstract.Theaimofthisstudywastoinvestigateanunderlyingmechanismoftheapparenttolerance of
Africanizedhoneybees(AHB)toVarroajacobsonimitesinMexico.Thiswasachievedbyconducting
thefirstdetailedstudyintothemitesreproductivebiologyinAHBworkercells.Thedatawasthen
compareddirectlywithasimilarstudypreviouslycarriedoutonEuropeanhoneybees(EHB)intheUK.
Atotalof1071singlyinfestedAHBworkercellswereanalyzedandcomparedwiththedatafrom908
singlyinfestedEHBworkercells.Therewasnosignificantdifferencebetweenthenumberofmother
mitesdyinginthecells(AHB52.0%,EHB51.8%);themeannumberofeggslaidpermite(AHB5
4.86,EHB54.93);thenumberofmitesproducingnooffspring(AHB512%,EHB59%);anddevelop
mentaltimesoftheoffspringinworkercellsofAHBandEHB.However,therewasamajordifference
betweenthepercentageofmothermitesproducingviableadultfemaleoffspring(AHB540%,EHB5
75%).Thiswascausedbytheincreasedrateofmiteoffspringmortalitysufferedbythefirst(male)and
second(female)offspringinAHBworkercells.Therefore,onlyanaverageof0.7viableadultfemale
offspringareproducedpermiteinAHB,comparedto1.0inEHB.
Keywords:Varroajacobsoni,Apismellifera,mitereproduction,Africanizedbees,mitetolerance

Introduction
ThemiteVarroajacobsoniOud.currentlyposesthelargestworldwidethreattoApis
mellifera L.honeybees.Theproblemismost acuteinregions withatemperate
climatewhichhavebeesofEuropeanorigin (DeJong,1990).However,intropical
regionssuchasBrazil,wherethebeeshaveanAfricanorigin(Africanizedbees)the
miteseffectonthecoloniesappearstobereducedtothepointwherenocontrol
measuresarenecessary (DeJongetal
.,1984;DeJong,1996).
IntheStateofYucatan,Mexico, V.jacobsoni wasfirstreportedinSeptember1994
andasurveyof603managedcoloniesin1996revealedanaverageinfestationlevelin
adultbeesof4.3% (Medina,1998).Sincethepercentageofafricanizationinthe

*Towhomcorrespondenceshouldbeaddressedat:12BurtonFieldsClose,StamfordBridge,York,
YO41LQ,England.email:s.martin@csl.gov.uk

660
survey area is high and increasing annually (QuezadaEuan et
al., 1996) the
situationinitiallylookedsimilartothatfoundinBrazil (Morettoetal
.,1995).
ThereasonswhyAfricanizedbees(AHB)intropicalregionsareabletotolerateV.
jacobsoni,unlikeEuropeanbees(EHB)havebeenattributedtothefollowingvarious
factors:

1.
2.
3.
4.
5.

Reducedmitefertility(reviewedin Martinetal
.,1997);
Smallerbroodcells (MessageandGonalves,1995);
Honeybeedevelopmentaltime (Moritz,1985);
Food(pollen)availability (Morettoetal
.,1997);
Increasedhygienicbehavior,viabroodremoval (CorreaMarquesandDeJong,
1998)andgrooming (Morettoetal
.,1993);
6. Climate (DeJongetal
.,1984;

Morettoetal
.,1991).
Althoughseveralofthereasonsgivenaboveinvolvethereproductiveabilitiesof
themites,adetailedstudyintothemitesreproductivebiologyinAHBcolonieshas
not yet been undertaken. Therefore, the aim of this study was to investigate an
underlyingmechanismofAHBtolerancebycollectingdataonthefertility,fecundity
andsubsequentoffspringmortalityofV.jacobsonimitesinfestingworkerbroodof
AfricanizedhoneybeecoloniesinMexico.Thedatacouldthenbecompareddirectly
with a similar study previously carried out on European honey bees in the UK
(Martin,1994).
Materialsandmethods
Ten Africanized bee colonies were established using mated queens captured from
swarmsandidentifiedmorphometricallyaccordingtothetechniquedevelopedby Daly
andBalling(1978), usingthecomputersoftwareAFUSDA7(R.Rubink,unpublished).
All coloniesbecamenaturallyinfestedin1995andreceivednotypeofmitecontrol
during the study. From February 1997 to October 1997 estimates of the comb area
occupiedbysealedworkerbrood,pollenandhoneyweremade.Inadditionmonthly
samplesofworkersealedbrood(8 3 8cmsection)werecollectedfromeachcolony
betweenFebruary1997andJanuary1998.Wherepossiblebroodwhichhadbeensealed
for longer than 230 hours (at least at the yellow thorax stage of development) was
collected.Thecombsectionswereimmediatelystoredat25Ctokillandpreserveany
mites.From100to150cellswereopenedineachsampleandthedevelopmentalstageof
thebeeandanymitespresentdetermined,usingtheontogenicdevelopmentchartsin
Martin(1994).Inadditionthefemaledeutonymphswereclassifiedinfivegroups(AE)
dependingontheirsize,usingthephotographsin Ifantidis(1983)asaguide.Thismakes
itpossibletoreconstructthemitefamilywithinacellandthendeterminethemortality
ratesoftheoffspring.Sincethedevelopmentofeachoffspringisseparatedbymorethan

a24hourperiod,itisclearifanyindividualoffspringismissing.Ifthiswasthecasea
closerlookwithinthecell

661
oronthebeepupainevitablyrevealedthedeadmissingmite.Onlyinfiveofthe
1,071reconstructedfamiliescouldthemalenotbeaccountedfor.
Priortostartingthestudybothauthorsspentseveraldaystogetherensuringthatall
themitesstagescouldbeidentifiedtothesamedegreeofaccuracyinbothAHBand
EHBcells.Alsotherulesusedtoreconstructmitefamiliesandtheclassificationofa
miteasliveordeadwhensampled,werediscussed.Thisensuredthatthedatafrom
the two studies was compatible and that differences would not arise from the
methodsofdatarecordingoranalysis.
Unlessotherwisestatedallanalyseswascarriedoutoncellswhichwereinvaded
byasinglemothermite.Tocomparethedevelopmenttimeofthemiteoffspringin
AHBworkercellstothatdeterminedinEHBcells (Martin,1994), thenumberof
each mite developmental stage found during each bee development stage was
recorded,i.e.duringthepinkeyedbeestage16and22ofthefirstmiteoffspring
were protonymphs and deutonymphs respectively. It was then assumed that the
changeofaprotonymphtoadeutonymphoccurred42%(16/[16122])throughthe
pinkeyedbeestage.Therefore,sincethepinkeyedstagelasts20hoursandstarts
140hoursaftercellsealing,thefirstdeutonymphoccursonaverageat1402203
.42 5 148hoursafterthecellissealed.Thesecomparisonsarepossiblesincethe
averagepostcappingperiodofAHB(275h,RosenkranzandEngels,1994;278h,
Vandame,1996)issimilartothatofEHBintheUKstudy(282h, Martin,1994).

Results
Ofthe11,655AHBworkercellsstudied1,946(17%)wereinfestedwithatotalof
2,347mothermites.Ofthese,1,071wereinvadedbyasinglemother.Thisdatawas
comparedwiththe908singlyinfestedEHBworkercellsreported inMartin(1994).
IntheAHBcolonies2.0%(40mites)oftheinvadingmitesdiedinthecellwhichis
similartothe1.8%foundinEHBstudy.Therewasalsonosignificantdifference(d
51.37)betweenthemeannumberofeggslaidbymitesinAHB(mean54.86,SD5
0.58, n 5 911) or EHB (mean 5 4.93, SD 5 0.54, n 5 131) worker cells and the
developmentaltimesoftheoffspring (Table1).Inadditionthenumberoffemalesnot
producinganyoffspring(nonreproducing)wassimilar(12%inAHB;9%inEHB)inthe
twostudies.
The major difference between the studies was the percentage of mothers
producingviableoffspring(atleastonematedadultfemaleoffspring).InAHBthis
falls to around 40% by the time the bee emerges, compared with 75% in EHB
workercells (Table2). Themajorreasonforthisdifferenceistheconsiderably
higher rate of mortality suffered by the first (male) and second (female) mite
offspring in AHB worker cells (Fig. 1). The level of male offspring mortality

remainedconsistentduringwingpadsgraytorestingstagesofthebeebrood
whileinalloftheotherstagesitincreasedwithtime (Table3).

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