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Departmental Bulletin Paper / 紀要論文
Comparative biology of the five Japanese
species of the genus
<special>Vespa</special> (Hymenoptera,
Vespidae)
日本産スズメバチ属 <special>Vespa</special>
(Hymenoptera, Vespidae) 5種の比較生態学的研究
Matsuura, Makoto
松浦, 誠
三重大學農學部學術報告 = The bulletin of the Faculty of Agriculture,
Mie University. 1984, 69, p. 1-131.
http://hdl.handle.net/10076/5564
ComparativebiologyorthenveJapanesespecie50rthegenus掩岬〝（Hymenoptera，Vespidae）
1
Comparativebiologyofthe蔦veJapanesespeciesofthe
genus物a（Hymenoptera，1砲spidae）
Makoto M＾TSUUR＾
CONTENTS
2
1）Queenne＄t
2）Changein shape、、・ith tIleemergenCeO（
WOrkers
3）Developednests
5▲Thermor喝ulation
6．Nestdefもnceandaggressiveness………………38
7．DisctlSSio！1
1一−一
3．Modiricalio】10r neSt Sile
ヰ．Relocation o＝1eSl
8ノ Qノ 0 −1 3 5 5
▲’1 1 1 1 1 2 2
2．Nest5truCture
7
1．Nest sites
7
1．Pre・neStlngperiod
2．Solitaryperjod
3．Cooperativeperiod
ヰ．Po）ycthicpcriod
5．Reproducliveperiod
6．Hibernatingperiod
Il．NESTING HABITS
′0
1‥へNNUAL CYCLE
2
lNTRODUCTION
METHODS
TAXONOMY AND DISTRIBUTION
38
1‖、FEEDING HABITS
1．Ⅰもod sources
1）Carbohydrates
ヱ）Protein
40
44
2．R）Oddependenceor物α汁叩JぐαOn
poiistine wasps
1）Co11eclion orpre〉・
2）Pheno10gyOrVisitto撤′ね′甜COlonies……53
3）Ratioor劫／如郎ノαdwなαぞCOlonies
attackedbyl鹿騨＝相画加……．…………‥54
ヰ）Predation period on a Po／istesncs【……‥．．56
5）Fateorattackedcolonyor撤〟∫∼郎………‥56
3．Grouppredationby物α∽α〝（ね油？ねon
nestsorhoneybeesandsocialwasps……．57
1）Visitsby掩呼びSPeCiestoapiaries…………57
ReccivedJune30．1984
2）Phenologyorattacktoapiaries……‥，…59
3）Phenologyorattack on othersocia）
vespid species
4）Predatoryha抽s
5）FoTagingdistanec
6）Ralioor＼・eSPincnesIs auacked and
destroyedby掩聯＝那胱加血紋‖…‥‖…65
7）Auackon hi＼・eS OrlheJapanese
honeyl）eちカタおぐem〃〝…‥………………‥66
4．Intra−andinterspecificrelationshipsat
わod source
l）DonliIlanCe rClationships amongquecns
Visitingtreesap
2）BehavioursofoverwirlteredqueerlSin
theel′enl一喝
68
68
71
3）1nterspeciricrぬtionshipsamoI鳩WOrkers
72
at fbod sources
74
5．Discussion
ⅠⅤ．DlVISlON OF LÅBOUR…………………………75
l．Beha＼，iouralchangeorthequcenin early
ncstingstageinrelalionto＼、・OrkercnlCrgCneC75
1）Extranidalactivities…………………………．75
−
ニ．∴∴
＿∴
1）Emergenceorworkers…‥‖…………………80
2）Behaviourorworkerssoona托eremergence80
3）Orientation nighland foragingtrips‥‥‥…80
4）Activitiesormarkedworkersduringtheir
adultlifも
3．Di＼・ision orlabourin an orphan colony”．．．．．8J
l）Be‡1訓iourorworkersandcolonydevelopme！1t84
2）DivisionoflabotlramOngWOrkers………‥85
3）Characteristicsorlayingworkers…………‥86
4．Discussion
1）Behaviouま・alchangeorqueeIlin early
nesting stage
2）Relationsbetwecnlabouranddayage
Makoto MATSUURA
2
89
3）Adulthornet5
3）DivisionoflabourinanorphancoIony‥‥89
4．Productionofsexuals……‥．…………………lO7
1）NumberofsexualsproducedpercoIony…107
V・POPULATIONDYNAMICS……‥，‥‥…．．．”．”．．90
1．Surviv（〉rShipcurvesof tb5PaWOrkers……．．…90
2）Larva：WOrkerratio…‥‥…‥…………‥‥．．．1O8
l）Lengthofde、・eloprnentalstage……．．．…．…．90 5．Nest survival
2）Sur、′i、・OrShipcurves foralldevelopmen【aL
l）Survivorshiporcolonies．…‥…．．．．”．．…．．．110
2）Mortalityfおtor5
射
3）Survivorshipcurvesforadultworkers……．92
6．Annualfluctuationsinthent】mberof
4）臨teorbroodatemergenceor5eXuals．…‥93
物αCOlonies
2・Seasonalchangesinthepopulation
7．Discussion
StruC【ureOflセspacolonies…．．‥‥‖‥……95
1）Survivorshipcurves．…”．…………‥．．．．．．”．．116
1）Numberofcells
95
2）Seasonalchangesinpopulation strueture‥117
2）ImmattlreStageS
97
3）Mortalityfactorsatco量onylevel‥‥‥‥‥…．1ま9
3）Adulthornets
101 VI．GENERAL DISCUSSION………………………120
3．Seasona）development ofasinglecolonyof
SUMMARY
陀岬〝J和〆Cα
105
ACKNOWLEDOEMヱNTS………………………125
l）CellconslruCtion
105
REFERENCES
2）Immaturesは酢S
105
摘要
INTROI）UCTION
ThesubfamilyVbsplnaeisdistributedmainlyinthenorthernhemisphere，Particularlyinthetemperate
ZOneS．Thissubfami1ycontainsonlyfourgenera，PIDl∫eSPq．掩5PLlk7，Do／ichol・e5PLL／aandIJ＞5Pa．LeavingPro−
VeSP￠aSrnallgenuswhichisrepresentedbythreespeciesfoundonlyinSoutheastAsia，theotherthreewere
PrOSperOuSgeneradevelopeddifferent modesoFlift，thoughwithaconsiderableo＼′erlap．Neither掩spzLIE7
norDolichove軍ujaisdistribtltedintropICalreglOnS，andtheyarecompletelyadaptedtotemperateclimates．
Thegenus物aismostlydistributedinthe王ndo−Malayanarea，butsomeofthespeciesarefbundtobe
dominantin temperate regions．
AccordingtoEDWARDS（1980）thegenus物ahas22speciesandofthese，SeVenSpeCiesaredistributed
asfarnorthasJapan（M，＼TSUUJuandYAM＾NE1984）．IntemperateEurope，thereisonlyonespecies，掩5Pa
C柑b7thWhichhasalsodispersedsecondarilytoNorthAmerica，Wherenoendemicspeciesoccurred．Incon−
trastto挽puねandDo／ichove軍uれthebionomicsoflセ軍ahavesofarbeenlesswellstudiedbothinEurope
andNorthAmerica，PrObablybecauseofthesmallnumberofspeciesandtheirhighlyaggressivedisposition，
despiteofthepioneerworksofJANET（i903）inFrance．Whilethegreateststridesinourunderstandingof
hornetbiologyhavebeenundertakenontheorientalhornet，陀5PaOrieJl（a／isinIsraelsinceBol）ENHEIトtER（1930，
1933），IsHÅY（1964−）andhiscolleagueshavenowmadeintensivestudiesontheethologyandphysioIogy
OFthisspecies−
1nJapan，Whichseemstoha＼′emOStrePreSentativesofthisgenusinthetemperateregions，man）・Ofthe
reportsonthelifeofthehornetsareonlygeneraldescriptionsandveryftwglVedetaileddata．
Since1957Ihavebeenengagedinbiosociologicalstudiesonthisgroupandhaveobtainedsomeinforma−
tiononthebionomicandethologlCalcharactersofthisgenusinJapan．Thepresentstudyintendstoap−
PrOaChthecomparativesocio−bioIogyofJapanese物ahornetsthroughextensiveecologicalresearchof
theirhabitsandlifehistory．Itishopedthatthisarticlewillprovideabasisfbrfuturecomparativestudies
notonlywithinthegenus，butalsoforcomparisonwithothersocialwasps．
METHOI）S
Studyarea
ThefieldworkinthepresentpaperwasmainlycarriedotltinandnearKibi−Cho，WakayarnaPre仁，SOuthern
Japan（Fig．1）．KibiisasmalltownalongtheRiverArita，Situatedca．20kmsouthofWakayamaCity，about
Comparati＼′Cbiologyorlheri、・eJapanesespecjesorthegenusI々spa（Hymenoptera，Vespidae）
3
′
■b
○′
．♂■ 旧n且順由江萱墓！些勺
・Q▲
Fig．1．ThemaportheJapan andthestudyarea．
Tbblel．Main climaticcondilionsin Kibi（1962−1975）
7ヒmperalure（OC）
m
（mnl）
7
7
0
1
7
つJ
Q′
7
エリ
5
7
ハブ
‘リ
′hリ
2
4
ーん
7
4
5 8
7 3
4 816名
∧U
7 4
16 12
Qノ 2 7 3
2
7 8 ︵y
27 21
17 12
9
3 7 つん 2
・・l ■・ま Z 2 ・・l l
■J
3
O
23
4
31
3
11．0
−5．4
︵j−7
a7
0
ーJ
27
30
0
Qノ
5
26
26
35．7
封
つJ
20．0
ぐJ−ヽ︸
23
Mean
Extremes
14
3．J
柑
∧U
13
￠U
9
5
9
﹂1
aX
m
O Jan勒Mar毎MayJu。Jul輌Se。OCt甑De。
㌣．mu
M
n
h
M
Rainra11
Tbtal18Z5．0
5
2
50−250mabovesealeveLMainclimaticconditionsinthelocalityaregiveninThblel．Theaヽ・erageClimatic
COnditionsare：meanairtemperature15．50C，thehottestmonthAugust（mean27．OOC，meanmaXimum3l．4OC）
andthecolde＄trnOnthJanuary（mean4．8OC，meanminimumO．70C），meanannualrainfal118之5．Omm，being
mostabundantinJune（238．9mm）andJuly（231．6mm）．Untilrecentlytheareahadbeencoveredwithsecon−
darymixedfbrestsmainlyconsistingofpines，deciduousoaks，Cherries，etC．，withsomeremnantsoftheprimary
lucidophylousforest，Shiiq，mmeHq，etC．Inrecentyearsthefbrestsarerapidlydiminishing，replacedbyrural
Or Suburban areas．
Theareaisremarkableinitsabundanceoflbpacolonies，andfivespecies，namely Vmandari17iajqpollicq
lくanaILsinsLJbris；Vcrvbroj7avq舟SCiatait（simi／／imaxunthppteruand VTrqPicapu／c／7rq，arefbundthroughout
their acti＼・eSeaSOn．The followingdescriptionsaregiven mainl）rbased upon these fivespecies．
Makoto MATSUURA
4
Melhod50−ObserY8tions
ThedataforthisstudyweregatheredmainlylnthefieldinandnearKibトchoduringtheperiodfrom
1964to1975．Duringthecourseofthisstudyextensiveobservationsinthefieldhavebeenmade，mOreOr
lesscontinuously，Onannuallifecycles，neStinghabits，fbodhabits，andactivitiesofadultsinsideandoutside
the nest．
Nestjnghabils：Manynestswere10Catedbyextensivesearchinthestudyarea，OrbyfollowlngqueenSand／or
WOrkersflyingfrombaitstationsorfoodsourcestothenestingsite．Somelocationsofnestswerereported
by10Calresidents．
Allthenestswerenumberedaccordingtotheorderoffindingofeachspecieseveryyear．Atotalof622
nestswereobservedinandnearthestudyareaoverthepasttwelveyears，andconsistedof288nestsof V
∫f川J／／山Jq1640rlてαJ柑毎800r H〔Ⅷ血Ⅵ560rトて〃化Jl（ねr由れand3ヰorレ：什0〆ぐ仇
Fbodhabi（S：Observationson food habitsweremadenot onlyin the Field butalso atthenestentrance，
Wherereturningfbragerswithpreywerecollectedtoexaminethespeciesofprey．Dataonphenologyofhornets
CapturedbytrapsattachedtohiveswerepreparednotonlyfromthesamplesinKibi，butfromthoseoffered
byseveralbeekeepersinAkita，1ぬmaguehi，FukuokaandNagasakiPrefectures．
TheFieldworkonthebeha＼′iourofintra−andinterspecificdominancerelationshipsamongpost−hibernalion
queensatfbodsourceswasearriedoutatdifftrentgrovesofQuerrustreesduringtheperiodfromApril
tolateJulybothin1966and1967．In1966theobservationsweremadeatagroveconsistingof7Que7℃uS
acutま∬imaand3Qgねucqwhichwere7∼10minhight，Ofwhich2treessecretedtreesap．In1967theobser−
Vationsweremadeatahillyelevation244mhigh，WherethevegetationwasacopseconsistingoflO￠
aculLssima．Inordertodistinguisheachindividual，まnbothyearsallthequeensof物athatvisitedthe
treeapertureweremarkedwithcolouroiトpaintsontheirmesonotum，aCCOrdingtotheorderoffindingof
eachspecieswhenshehadalightedonthetrunknearbythetreeaperture．Observationsweremadeatas
nearaplaceaspossibleandintheevenlnganelectrictorchwasusedtolightthehornets．
QIL〝／i（atit・eandqlLa［1（i（ativechallgeSq［qlLeenaC（ivi（ies：Thestud）・WaSCarriedoutmainlyonanincipient
nestoflてcrabro（C6601）attachedtotheceilingoftheatticofastorehousein Shimotsu−Cho，Wakayama
Pref．in1966．The nest was situated about50crn ftom thewindow andindividuals ofthenest hadto take
arouteonfbot37cmalongthewa11and50crnalongtheceilingtoreachthenest．Observationsweremade
fbr28days（120hours）duringtheperiodfromJune2ndtoJuly28thin1966atirregularintervals．Atthe
lastobservationonJuly27th，thenestconsistedoftwocombsand190cellswithaqueenand15workers，
WhichwerekilledbyinsecticidesprayedaroundthenestonJuly28th．
RelationsbetweenlabouranddqyageirlWOT・keTT：Thestudywascarriedoutwithqueen−rightcoloniesof
Fi＼′eSpeCies，V：simi／Iima，Il：anaIi51T／：crabrQ．IlmalZdariniaandIて（rOPica．New）yemergedworkersineach
SpeCieswereglVenanindividualmarkontheirthoraxwithoil−paintsonthedayofemergence．lnfburspecies，
t（TrOPica，Ilana／hlt／：cmbroand Vmandarinia，ne＼Vlyemergedl＼′Orkel■SOnthecombs≠′ereCarefullymark−
edwithoutpickingthemupfromthecombsurface．王n Vsimil［imaitisverydifficulttomarkthethorax
directlyofnewlyemergedworkersonthecombswithoutanydi＄turbancebecausetheyquickiyirritated．
Therefore，ltookworkerswithinabout10hoursafteremergence汁omanothercolony（S6802）ofthesame
SpeCieswhichwaskeptinanothernestbox，andreleasedthemintotheobservationnestafterrnarking．The＄e
teneraiworkerswerealwaysacceptedwithinonedaywithoutanyslgnSOfhostiiity，andtheyimmediately
joined theindigenous＼VOrkersinperformingany tasksin thenest．
Thetotalnumberofindividual）ymarkedⅥ・Orkerswas57inIJ：sirTll／［ima．35inレ：aJJaIis，39inIイITla／卜
darinia，36in VtTPPicaand80in Vcmb7ⅥreSpeCtively，anda11ofthemwerefb1lowedtoestimatenotonly
thedayofthefirstfbragingbutalsothelongevltyOfworkers．Ofthesemarkedworkers，autheactivitie
WereObservedonsomeindividualsduringthciradultlifbatirregularintervalsinthedaytimeandrecorded
togetherwiththedurationspentineachactivitywithintheerrorofabout j＝3seconds．Thenestsobserved
are presentedin the fotlo≠・ing notes（P．5）．
DivLsiondh7bourinofPhancolonies：Observationsonorphancoloniesweremainlymadeonanestof V
analis（A6705），WhichwascollectedinNachikatsuurarcho，勒kayamaPref．onJuly23rd，1967．Thecolony
ComparativebiologyortheriveJapanesespeciesorthegenus掩叩α（Hymenoptera，Ⅶspidae）
Species
NesとObserved
No．orworkers
Observed
1ndividualNo．
∫∫′”′／伽7α
S6801
3
Nos．8，9，10
α〃α〆ね
A7ユ07
′”α〃血r∼〃′α
如月Jぐα
ぐ柑∂柑
M7401
T6803
C6601
2
Nos．3，4
5
2
No5．5，6
Nos．2，3
4
Nos．38，39，
40，ヰ1
12：00
Time or
emergenCe
Tota】time or
observation
−16：00
July 12th
1986
25days
（102h）
12：10．
1ユ：50
August 3rd
197ユ
28 days
（82h）
8：20，
12：20
July28ぬ
197▲1
27 days
（81b）
12：00
−15：00
8：30
−16：00
July25th
1968
Juiy2lst
1966
60 days
28 days
（140b）
（70h）
Withallnestmembers（1queenand16workers）wasintroducedintoanobservationboxwhichwascon−
nectedwiththeexteriorbymeansofabambootube．Theq11eenWaSremOVedartificiallyonthedayofcol−
lectingandreturnedtotheorlglnalnesttheseconddayafterremoval．Butshewasimmediateiyattacked
andkilledbytheworkers．AutheworkersattimeofcollectlngthenestweremarkedwithcolouroiトpalntS
Ontheirmesonotum．Atotaiof72hours（50days）ofobservationwasmadeonthebehaviourofthe10
markedworkersduringtheperiodfromJuly23rdtoScptember15th．
SurvivorshわqfcoねnymembeFT：Inpreparinglifetablessurvivaldataforbroodandadultsoffive Ve軍a
SpeCies．i．e．，SimiI／ima，ana［is，mandarinia，（rOPicaand crobro，＼＼′ereObtainedfroTndi汀erentsources，and
COmbinedintooneduetosometechnicaldifficultiesinfol10Wlngthewholelifehistoryofparticularin−
dividuals from eggto adult stage．
Observationsineachspeciesweremadepr亘nclpa11yonthecolonieslistedearlier．But，in n′rq錘〝the
datafromanest（T7401），WhichIobservedduringtheentireperiodfromnestfoundationtodisintegration
areused．Inordertodeterminethelengthofeachdevelopmentalstageanditssurvivalin5uCCeSSiveimr
maturestagestheywcredividedintothreestages；eggS，1arvaeineachinstarandcocoons．Eachbroodcell
WaSdailymappedandphotographedthroughouttheobservationperiods．Ceumapsweremadeonceaday，
andthedatesofoviposition，hatching，COCOOnSpinningandemergenceofindividuaisofthebroodwere
estirnatedfromthem．Theseeventswereregardedtohaveoccurredonthedaywhen蔦rstrecorded．The
SurVivalofbroodduringthecocoonperiodcouldnotbeobserveddirectlybecauseofitstechnicaldifficulty，
andlarvaeorpupaewithincocooncellwereregardedasalivewhenthecellcapsremainednormal．The
deathofbroodwasalsoascertainedbytheemptycellsinbroodcircles，becausethebroodinacombexist
inconcentriccirclesoreggs，lar、・aeandcocoons，andblanksinbroodringsindicate＼＼・heredeathhasoccur−
red（SpRADBERY1973）．
Allthenewlyemergedworkerswerenotobserveddirectlyatthetimeofemergenceandsuccessfulemergence
WaSCOnfirmedbytheexistenceofteneralworkerswhichusuallyinsertedtheirbodyintotheemptycellfrom
Whichtheyhadjustemerged．Inordertoestirnatethelongevityofworkers，neWlyemergedworkerswere
glVenanindividualmarkonthemesonotumwithpaintsonthedayofernergence．Thetotalnumberofin−
dividua11ymarkedworkersineachspeeieshasalreadybeenmentioned．Eachindividualwascheckedfrom
daytodayandtheexaminationwasrnadeprlnCIPa11yeveryevenlngaftertheirflyingactivityhadceased，
withoccasionalirlSpeCtioninthedaytime，untilthedisappcaranceofallmarkedworkersineachcoIony．
Theworker’slastdayalivewasassumedfromthedatewheneachindividualdidnotreturntothenestany
more，beeause＼VOrkers almost never diedin the nest．
SeasonL7／changeinpqpLL（a（io11S（nlC（LJre：1nordertocstimatetheseasonalchangesinthepopulationstruc−
tureofeachspecies‡col王ectedatotalof393nestsinaustagesofdevelopmentfrom1964to1975，Which
COnSis【ed of150nests of V．simi／／ima，Yart／hop（era，122nestsofl′．L711aIisi／JSLJIaris，38nests of V．crabro
Makoto MÅTSUURA
navo払sciata，49nestsorH椚α〃血r玩∫αノ叩0〃∼cαand34nestsorH汁叩JC叩〟ね烏m．刃loftbenestsexcept
largecoloniesofレてsimiI／tmaweretakenwithreturningworkers，Whichwerecapturedwhenthenest＼VaS
revisitedaboutoneortwohourslaterinthedaytime．Anumberof Vsimi〟imanestsatthelaterstageof
nestingwereoccasionallytakenintheevenlngWiththehelpofchoIoroformanda11anesthetizedworkers
insideorontheenvelopewerecollectedtoexaminethecompletecontents．Thenumberofcombs，Ce11s，Pi11ars．
immaturestagesandadultswerecountedineachnestonthedayofcoliecting．
Developmentofasinglecolony＼＼′aSalsoobservedonanestoflてJrOPica（T7401）which＼＼・aSfoundedin
awoodenboxatⅥ血kayamaFruitT†eeExperimentalStationinKibionJune9th，1974．Changesinthecol−
Onydevelopmentandpopulationstructureofthenestwerefbllowedatitsnaturaisiteduringtheactiveperiod
from neslfoundation to disintegration onOctober15th，1974．
Nestsurviva］：DuringtheobservationperiodIfbundatotalof59nestsofVana胎insuJarisatanincIPlent
Stage．Thesenestswerecensusedatirregularintervalsinthefielduntiltheendofeachnest．Thedatafrom
manyunsuccessfulnestsofeachspeciesfoundinthefield，tOgetherwithrnysuccessiveobservationsonthe
activenests，gaVeSOmeinfbrmationtoestimatethesurvivalof挽甲aCOIoniesandvariousfactorscontributing
tocolony termination．
加〃〟αJβ〟C加地乃げcoわ野〃〟椚∂椚：Therecords紺emainlybaseduponthematurenestsorthe賞vespecies，
IくsimiIIima．V：ana／isIV：m7brq VmandarEniaandレ：TrqPica，Whichhavebeenobservedinandnearthe
SurVeyareaduringa12−yearperiodfrom1964to1975．Besides，aSyStematicsearcbfbrallcoloniesof
SimiHimGinthestudyareawasmadeannuallyinlateautumn，Whenthecolonieshadaimostcompletedtheir
nests，duringtheperiodfrom1964to1975．Thearea，COVering300hectaresconsistsoffoursub・areaS，Mu−
tsugawainKanayかCho（Ca．75ha），1ゐshikawainXanaya−Cho（Ca．80ha），1bsbimiinXibトcho（Ca．50ha），
and協madain％asかCho（Ca．95ha），auOfwhicharevillagesamongthehi11sofabout50−500metersinheight．
TAXONOMY AND DISTRIBUTtON
ThesuperfhmilyVbspoideaisdividedintothreefamilies；Masaridae，EumenidaeandVtspidae（RrcHARDS
l粥2，1972）鯵Thesocialwasp50rVespidae払11intothreesubねmilies；Stenogastrinae，PolistinaeandⅥさSpinae．
AccordingtohismostrecentrevisionortheVとspinae，EDWARDS（1980）recognizedねurgenericnames：伽−
γ朗pαA5hmead，掩甲〟ぬThomsom，か0眈如v叩〟ねRohwerand掩鱒αLinnaeus．Thevesplnegeneraare
primarilyatropicalAsiangroupandhavepenetratedintothetemperatereglOnSOfEurasiaandNorthAmerica．
Auve＄plneSareeuSOCialorsocialparasitesontheireusocialrelativesandarenoteworthyfbrtheadvanced
StateOfsocialityrelativeto mostofthePolistinae．
Thegenus挽spaisdistinguishedfromotherVespinaeprimaril）・byitslargesizeandtheexpandedcondi−
tionoftherearpartofthehead（BEQUAERT1930，1931，DuNCAN1939，VECHT1936，1957）．Theoriginaldescrip−
tiono‖句卯madebyLJNN＾EUS（1758，SystemaNature）comprisedallthethenknownspeCiesno＼l∫included
inthesubfamily．Thegenus t／bspahasbeenre＼′isedb〉・S＾USSURE（1853−1858），THONSON（1869），BuYSSON
（1903−1905），BEQUAERT（1930−1939）andVECHT（1959），andaspresentlydelineateditcontainsapproximate−
1y22specieswhicharemainlycenteredinsoutheasternAsia．Onlytwospecies，VcTUbfVand Vorientalis，
arewidelydistributedinthewesternhaifofthePalaearcticreglOn．
1nJapanthefbllowingformshavesofarbeenrecorded（MÅTSUURAandYAM＾NE1984）：
Forms
（Japanese name）
l．t！卿nts仰／7isLINN＾EUS1764
Distribution．Ryukyuislands．
（恥umaguro−SuZumebachi）
2．V：ana／Lsli（1Su／arLsD＾LL＾ToRRE1894
（Kogata−SuZumebachi）
Distribution．The≠′holecountry．
3．打ご用か柑ガ〝V（押∫C如αCÅMERON1903
（Mon−SuZumebachi）
Distribution．The＼＼血olecountry．
4．巨め′ねw∫如∼AND鼠E1884
（Chairo−SuZumebachi）
Comparativebio10gyOrtheriveJap8neSeSpeCiesorthegenus掩岬〃（Hymenoptera，Ⅶspidae）
Disこribu貢on・Hokkaido，Honshu（Central）．
5．H沼α〃（ねr血由加0乃加RÅDOSZKOWSK王1857
7
（ら一SuZumebachi）
Distribution．The、、・・holecoutltry．
（Kebuka−SuZumebachi）
6．ト：∫山JJ仙川〟∫／川／／／山J〝SⅣ‖T↑11968
Distribution．Hokkaido．
7．H∫′J乃′／／f椚αズα〃′如卿g柑CÅMERON1903
（Kiiro−SuZurnebachi）
Distribution．Honshu，Shikoku，Kyushu．
8．巨汗qクJCαがα舟〟SoNAN1935
Distribution．T§ushimaislan（董．
9．ト：什0〝血／00C／JOOビ〃∫豆Bl三qし】八lミRl1936
（Esaki−SuZumebachi）
（Okinawahime−SuZumebachi）
Distribution．Ryukyuislands．
10．卜：什Op血plJ／c／ImBuYSSON1905
（Hime−SuZumebachi）
Distribution．Honshu，Shikoku，Kyushu．
AIsol≠・O rOrtllS，t／：biぐ0／orF′＼u火lぐ1US1787and r棚伽r∼BuYSSON1905，arereCOrded汁omJapan，The
formerspeciesrecordedbyBuYSSON（VECHT1957）inalIprobabilitydoesnotoccurinJapanandthelatter
recordedfromCentralJapanbyVECHT（1959）isunknowntomeasIhavethusfarnotencounteredtheform
COincidingwiththeoriginaldescription（BuYSSON1905）．
Ⅰ．ANNUAI．CYCI．E
Intemperateregions掩5PaCOloniesareannualandthenewcoIoniesarealwaysbeguninthespringby
aslngiefbcundatedqueen・Theannuallifbcycleof掩5PaaSpreSentedsubsequentlyinthischapterconsists
Ofthefollowingsixperiods（Fig・2），thoughalittlevariationindetailsofthecycleoccursamongthespecies：
l）Pre−rleSting，2）Solitary，3）Cooperative，4）Polyethic，5）Reproductive，6）Hibernating．Thesequencedoes
notessentiallydifferfromthatinothertemperateeusocialinsectswithannualco10nies；挽軍uJa，Bombus
and，Withsomedeviations，Aフ触e∫andeusocialhalictinebees・AbrierresumeisglVenhere．
1−1一（）−neSしinq
PERIOD
Solまヒary
Cooperati．vQ
poly（≧ヒ11まc
R愈prOducヒive
鵬b（≧rnaヒing
済［コ皿孤田巨ヨ［≡］
＞．tropICa
V．∂n81is
V c「8bro
V 什1こ1nd亡1rl11ia
V二 Siml＝ima
Fig・2・Lifehistoryoffjve物aspeciesinsouthwesteFnpartSOfHonshtl，Japan．
l．Pre−neSlingperiod
Thi5period，tObederinedexactly，meanSadurationfromthetime、沌enthequeensemergeftomtheim−
mobilizedstateofdiapausetothetimewhentheydecidethesitesforconstructlngtheirnewnests．Thedates
OfthefirstencounterofhibernatedqueensinthelastlOyearsarepresentedin′払ble2，Whichshowsthat
thetimeofappearanceisdifferentineachspecies．Judgingfromlもble2，tOgetherwithotherobservations
Makoto MATSUURA
8
Thblc2．Dateofthe nrst encounterofhibernated qucensofレ旨5PaSPeeies jn and near Kibi．
「ぬar
′る6
′68
′67
′69
′70
り1
ワ2
り3
ワ4
′75
Apr．12 Apr．5 Apr．10 Apr．29 Apr．15 Apr．20 Apr．17 Apr．20 Apr．14 Apr．18
r．椚α〃（ねr飯fα ‖ 8 ′′ 22 り 6 ‖11 ‖ 2（i ‖12 ’′ 24 ‖ 30 ′′18 り 27
V．simi／lilTlO
5
y
a′
つん
′
2
M
︵l一六Y
nけqクJぐα
‖ 28 ‖ 25 May2 ‖ 21May5 May3 ‖ 24 May2 ’′ 25 May2
つJ
r．α〃♂Jね
MayJun
y．ぐrα∼汀0
‖ 2 May9 ’′ 3
′’5
‖ 2
Jun．4 Jun．8
byme，Vsimil［imafirstcameoutoftheirhibernaculaeachyearinearlytolateApril．Both V：mandari77ia
andIてcmbzvgenerallyappearedfrommidApriltoearlyMay，Vana［tsinearlyMayand Vlrqpicaatand
aftertheendofMay．Thus，theinterspecificorderofspringappearanceis，Simi［［ima−marldari71ia−C／甘bro
一肌祓ぃ抒叩加（MATSUURÅ1971）．
Amongthesespeciesレ：si（T7i／／imaoccasionallyvisitsflo、、′erSOfCame／iajoponica，butotherspeciesseldorn
frequentnowers．Themos（importantfoodsourceofpost−hibernatingqueensisthesapofQELe汀ILStreeS，
Whicharepartic11larlypreferredby VmandarinjqwiththepeakofvisitsfromlateApriltoearlyMayas
ShowninFig．3．ThequeenstaketreesapnotonlyduringthedaytirnebutalsointheevenlngfbrseveraI
S﹂く﹁一凸−＞−□Z﹂
‖汽︶
T道心ち⊃N
Fig．3．Dailychangeofthenumberofo＼′erWinteredト卸amandqri（Liaqueens＼′isitinglreeSaPintheeYening
at acorpse，COnSistingofabout201reeSOr QLLerCJJ∫SPP・，in Kibi−Cho・
daysafterhibernation．Adistinctlineardominancerelationshipisrecognizedamongthesequeensvisiting
treesapandtbeintra−andinterspecificrelationshipswillbedi5CuSSedingreaterdetaillater（Chap．ⅠII．Fteding
habits）．Ithasneverbeenobservedtbatthe叩eCiesof物αCatChinsectsfbrfoodbefbrefoundingtheir
nests，inspiteofmycarefulobservations．
Itisrecognizedthatthefreshweightofpost−hibernatingqueensof物aisverymuchlowerthanthat
ComparativebiologyofthefiveJapanesespeciesofthegenus挽甲q（Hymenoptera，Vbspidae）
9
Ofthepre−hibernatingones．In VmandariT2iathemeann・eShweightofqueenswas3．46土S．d．0．21gjustbefbre
hibernation，andthat of34queens co11ectedin thesprlng Ofboth1966and1967was2．14土＄．d．0．22g
（MATSUURA1966）．
Itseemslikelythatsucharemarkabledecreaseoffreshweightiscloselyrelatedtotheirconsumptionof
fatbodyduring，andsoonafter；hibernation．Thehibernatmgqueensretainagoodquantityofねtbody
COVeringtheovarywithintheirabdomen．Theamountoffatbodyisalreadysmallsoonafterdeparturefrom
hibernaculaanddecreasesrapidlywiththecornmencernentoftheiractivities．AccordingtoSpRÅDBERY（1966），
fatcontentasapercentageofthedryweightinqueensof陀軍uぬspeciesisover40percent，Ofwhich30
PerCentisutilizedduringhibernation．Thequantityoffatcontainedinpost−hibernatingIゑ野aqueenSnO
longerincreasesinspiteoftheirfeedingactivityandlittlefatbodyisobservedafterMay（MATSUURA1966）．
2．Solitaryperiod
AllspeCiesof掩軍afbundtheircolonieswithasinglequeen，beingregardedashaplometrotic．Untilthe
emergenceofthefirstworkerithequeenengagesina11duties，includingnestfoundation，OVlpOSitionand
theprogressiveprovisioningtothede＼′elopinglar、・ae，aSis（ypicalinmostsolitarywasps．
Thequeenssearehfbrnestingsitesafterspendingtimeintakingtheirownnutrimentforashortperiod．
Thefirstobservationofsearchingfbrnestingsitesin Vsimi］limawasonMay4th，1965，andthelastobseト
VationonMay24th，1966andMay19th，1967．In Vmandari77iawhichbuildsundergroundnests，10bserved
firstonMay22nd，1967andMay19th，1968，aqueenrePeatedlyenteringsomeready−madeburrowsexcavated
bysnakesorsmallrodents，andjudgingfromtheirbehaviour，theyweresearchingfbrnestingsites．Although
lhaveFe、VreCOrdsofthedatesoftheotherJapanesespecies，aneStimationmaybegivenonthebasisof
thedatesoffirstspringappearanceofpost−hibernatingqueens（1もble2）andthedataonnestcontentsat
anearlystage（7hble8）：Vc用bmafterearlyMay，Vanalisaftermid・May，andIてtTqPicaafterlateMay．
IngeneralJapanese物aspeciesstartnestbuildinglaterthantheotherJapanesesocialwaspssuchas劫ぬtes
劫明卿研油and噸〟ね（MATSUURA1975）．
‡tmayperhapstakeaqueenofeachspeciesseveraidays，eVenWeeks，tOfindasuitablenestingsite．As
SOOnaSShechoosesadefinitenestsiteshestartsnestingactivity．Theprmclpalbuildingmateria王saregenerally
WOOdfibresmixedwithsalivarysecretion．Thefibresusedarederivedれ）mthecortexoflivingtrees，decayed
WOOdandrottenpartso‖ivingtrees．Thefirststepofnestconstructionistomakeapediceltosupport
asmaucomb．Afterthefirstcombconsistingoftwotofourcellsisbuilt，thequeenmakesanenvelopeto
COVerit．0＼′ipositionoccursalmostparallel、Vithcellconstructionandthe firsteggislaidassoonasthe
Firstcelliscompleted．
Asinglequeenofeachspeciesconstructsinaverage35∼40ceusandrearslO−15larvaeuptothepupal
Stagebyherself．ThesolitaryperiodlastsfbrabouttwomonthsfromiatesprlngtOearlysummerastotal
developmenttimesfromeggtoadultinthefirstbroodrequiresmorethan50days．Thenestinghabitsin
thisperiodwillbediscussedingreaterdetailbelow（Chap．ⅠⅠ．Nestinghabits）．
3．Cooperativeperiod
Asshowninlもble3，firstworkersof物aemergedfromJunetoJulyinthefollowingordcr：SimHlima
−Crabro−ana／is−mandari／血−（roPica．Thedifferenceinpeakdates＼VaS5to6weeksbetweenレ：slmi／／
and［く（rppica，thoughthetimeofemergenee＼′ariedamongcoloniesofthesamespeciesITheseworkers
arealwayssmallerinsizethanthoseemerginglaterbecausecelisproducedbyqueensarethesma11erthan
anyproducedlaterin theseason（lbble7）・
Thenewworkersbegintoengageavarietyofintranidalactivitiesasearlyastheirseconddayandperform
fbraglngtripsat2to4daysold，aSSOOnaStheyareproperlyendowedwithflyin＄ability．Aftertheemergence
ofworkers，thequeensarestillconcernedwithextranidalandintranidalactivitiesforawhile，thoughthe
behaviourofqueensvariesinbothquantityandqual辻yintbecourseoでcolonydevelopment．Thisperiod
isca11edthecooperativeperiod．Inthisstagequeensfirstceasetheirextranidalactivities，thenintranidal
activitiesexceptforoviposition．Theprocesswillbediscussedlaterindetail（Chap．VI．Divisionoflabour）・
Itseemslikelythatthelarg餅thetypicalcolonysizethesbo沈erthecooperativeperiodla摘・‡n㌣汁q錘偽
10
Makotoトl人TSUUR入
lbble3．Dateorthe rirst appearenceor拍spaworkersin and nearlこibi．
1漁ar
−66
r．∫J川J／／J〃！α
′69
り0
′71
り2
り3
り4
′75
Jun．19 Jun．14 Jun．10 Jun．8 Jun．12 Jun．9 Jun．9 Jun．11Jun．14 Jun．10
′′ 2＄ ’′ 25
‖ 28 ’′ 24
り 25
り 27
り 27 ” 21
2
0
つ︼
2
つ一
つん
0ノ
4
4
り‘
20 つ︼
つJ
つん
つー
3
3
▲JJ
2
1
つふ
4
2
′nV
O
2
3
9
ハV
Iノ．川α〝dα′／／l由
‖ 22
︵U
つ︼
′′ 22
y．〝娘偵
Jul．24 Jul．ユO Jul．24 Jul．25 Jul．23 Jul．2＝uL26 Jul．24 Ju）．15 Jul．20
つん
一′，什q桝Cα
’68
4
l′．仁和わ′O
’67
WhichhasthesmallestcolonysizcamongJapanese佗spaspecies，lhequeensparti亡ipatedinintranidaltasks
throughouttheirlife．Thequeensof Vana勧whichalsobelongtoasmallcolonytype（Fig．65），COntinued
toengageininternalactivitiesforabout7to8weeksaftertheemergenceofworkers．Ontheotherhand，
inthecaseofspeciesoftherelativelylargecolonytype，thequeensgaveuptheir“workertasks”after6
to7weeksinlイcmbroand2to4weeksinIてITlandarinEa．Andin somecolonies of V：simi／／ima，Which
Oftenhasmorethantenthousandceus，thequeensstoppedtheirintranidalactivities2to4weeksafterthe
eme柑enCeOfworkers．
Incontrastwiththenextpoiyethicperiod，thecolonyinthiscooperativeperiodischaracterizedbysIow
development andasmallnumberofworkers as shownin Figs．65−76．
4．Polyethicperiod
Asthenurnberofworkersincreases，thequeensofallspeciesexceptfbrthoseofVtrqpicabeeomerestricted
tooviposition，1eavingworkersalltheotherduties．Withtheattainmentofdivisiono‖abourbetweenqueen
andworkers，thelifbof物ashiftsfrornthecooperativeperiodtothepolyethicperiod，andthecoIony
undergoesarapidincreaseinpopulation．Thisexpansivestagelastsuntilthetimewhenmale＄andnewqueens
emerge，
Tbwardsthemiddleofthisperiod，COloniesofboth V：similfimaand Vm7bTVOftenrelocatetheirnests
fromtheirorlglnainestsitesinsmallspacestonew，mOreSPaCioussites．ThisstrategylSreStrictedtothe
twospeciesmentionedandadetailedaccountofthisremarkablefeatisgiveninthenextchapter．
Inthecourseofthisperiod物aworkersstarttodisplayobviousattraetiontotheまrqueen．Theycluster
aroundthequeenonthecomb，Orientatingtowardstheheadandthorax（Fig．4）．Someofthemcontinuauy
antennateandlickherbody，Whichisverysimilartotheroyalcourtinhoneybees．Thephenomenonisex−
Plainedbythepresenceofaqueensubstance，Whichhasbeenchemicallyextractedfromthequeenof V
Orienta］LgbyIsH＾Yetal．（1965）．ThisbehaviourisnotseeninthecoloniesofVtrqpicqwhichhasthesmallest
colonysIZeamOmgJapanese掩騨αSpeCies．AttheendoftheseasonlVOrkersが巨cJVみ和StarttObitetheir
queenfrequentlyandevencauseherdeath．Butthesituationinthreeotherspecies，Vana勧Vsimi［［ima
and Vmandariniqissomewhatdifferent，fbrtherehavebeennoobservationsofworkersiickingthequeen
Violently．Queensof挽汐a（excepting Vtrqpica）arecharacterizedbyahairlessandpolishedbodyinthis
period（MATSUURA1968），Whichmaybebroughtaboutbythelickingofworkers．
Astheendofthisperiodapproaches，theworkersofeachspeciesceasetobuildsmallcellstorearworkers
andbegintobuildce11sofalargersizetoproducesexuals，thoughthechangeinthesizeofcellsappears
totakeplacesomewhatgradua11y．AsthelargecellsinautheJapanese ya5PaSpeCies（exceptfbr Vtrqpica）
arealwaysbuiltsoonaftertheappearanceoraroyalcourtorworkersinthecolonyバheinitiationofla曙e
queence11sappearstobecontro11edbythequeenpheromoneasalreadypointedoutin Vorienta胎byIKÅN
etal．（1969）．ln Vt7Wic払WOrkersareapparentlyunaffbctedwhenclosetotheirqueenandnotgatherround
heちWhichsuggeststbatqueencontrolortheworkersi5doneinotherways．Atanyrate，tbel）rOOdofsexuals
in‡セ軍aCOIoniesarerearedduringthetirnewhentheworkerpopulationisatitspeak，aSdiscussedlater
in chap．V．Populationdynamics．
Thedurationofthepolyethicperiodseemstobecloselyrelatedtothelengthofcolonylifbineachspecies，
ComparativebiologyofthefiveJaparleSeSPeCiesofthegenusI句卿（Hymenoptera，Vespidae）
1l
Fig．4．Royalcourtinlセ3PaSPeCies：A，Fてcrobro；B，［・：aTla［is．
insuchawaythatthelongertheperiod，thelatertheemergenceoぎsexualsocぐurS．Theperiodlasts3∼3．5
monthsinthemostion針IivedcoIoniesofboth Vmanddriniaand Vsimil［imq2−2．5monthsinthemoderate
eolonies ofレ：ana／iもandl．5−2monthsin theshorト1ived colonies oflくc（t7bro．
5．Reproductiveperiod
Thepolyethieperiodcomestoanendwiththedeathofthefoundingqueen，Whichu＄uauyOCCurS3uSt
befbreorsoonaftertheemergenceofsexuals．Thereproductiveperiodlastsaslongasworkerh6rnetsare
StilL ative to feed the new sexuals．
Recordsorthesurvivalofねundingqueensobservedinfield−COllectednestsoreachspeciesareshown
12
Makoto M▲TSUURA
Thble4．Latestdatefoundingqueenswereobservedin rield−COllected nestsof Lセspaspeciesinand nearKibi．
Y毎払r
′66
′6？
′68
′69
′70
り2
り1′
′74
ワ3
′75
y，ざ如∫J才知α
Oct．20 0ct．12 Nov．3 Nov．11 Nov．7 0ct．27 Sep．27 0ct．10 0c【．23 Sep．3
V．crabro
Sep．22 Sep．22 0cし19 0ct．15
V．（川a／is
＿
′′ 20 Nov．1
y．椚α〃（ねrf〃ぬ
V．tropicq
‥17 Sep．17 ’’22
＿
＿
＿
＿
Sep．30 ‖19 ‥ 3
‥17
‥ 24 ’’13 Sep．1 Sep1110ct16
＿
Oct．18 0ct．10 ’′ 24
Aug．30 Sep．25 Aug．29 Sep．18 Aug．27
＿
＿
’′ 7
＿
Aug・26
Fig．5．Newqueensof Ve5PqCrqbro：＾．restinginthecells；B，rCStingontheenvclope・
T
ComparativebiologyorthefiveJapanesespeciesofthegenus掩岬α（Hymenoptera，ⅥきSPidae）
13
inTable4．Thequeensof7itrpptcqashort−CyClespecies，diedtowardslateAuBuSt，Whereasthoseofboth
Vmandarinjaand Vsimillimqlon針CyClespecies，SurVivedeventoNovernber．Theinterspecificorderof
disappearanceoffoundingqueensis，（ropica−CrObroqo（7aIis−mOndorinia−Simi］Iima，lVhichisinaccor−
dance＼＼・ith theorderofemergenceofne≠・queenS．Thelifespanofqueensin each species appearStO be
abou【Oneyear．
Withthedeathofafbundingqueenthecolonyentersadisorganizedstate．Sorneworkersfightoneanother，
andnestdevelopmentceases，thoughforaglngaCtivityandbroodnursingcontinue．After3∼4days，thecol−
OnyreCOVerSthelostharmonylnparallelwiththeappearanceoflayingworkers．Adetailedaccountisgiven
in ehap．1V．DiYision oflabou［■．
Themalesandnewqueensemergeatthesametime，Orthefbrmeralittleearlierthanthelattertowards
thedisorganizedstageoftheseasonalcycle（Cf．chap．VIIl．Populationdynamics）．Withtheappearanceof
SeXualstheexpansionoftheneststructureisstoppedorconsiderablyreduced．SexualsremainlOto20days
Withinthenest，andspendmostoftheirtimethrustingheadandabdomenintoemptyoregg−COntainlng
Ce11sattheeentreofthecomb（Fig．5）except foroccasionalfoodintakefromworkers（Fig．6）andlarvae．
Fig．6．1句卯trOPicanewqueenssolicitingaworkerforsali、・arySeereIion．
TheyhavenothingtodowiththedivisionoflabourinthecoIony．Atthelaterstageofthisperiodforaglng
activityofworkerschangesprinclpallyfromproteinfoodtocarbohydratessuchastreesap，fruitJulCe，etC．，
tofeedtheadultmalesandnewqueens．Thisresultsinadecreaseinattentiontothelarvae，Whichfaiito
befedandremaininthesameinstarforseveral＼＼′eeksbeforefinallyd〉・ing．Thedeathofthelastsurvi＼′ing
workersbringsaboutthecompletedisintegrationofthecolony．Thelastsexualstoemergecannotobtain
Sufficientfbod．Theyleavethenestwithoutmaturingandaudieofhungeraroundthenest．Ontheother
hand，餌11ymatureds誤ualswhosebodycolourhasintensiriedandbodyweighthasincreased（e・g・，鉦om
2．9gto3．5gin7（mandarinia，M＾TSUURÅ1966）departfromtheirnests．Departingsexualsdonotmakean
OrienTation flightand ne＼′erreturn tO theirnesl．
ThedisintegrationofcoloniesordinarilytakesplacefromearlySeptembertolateNovemberinthefollow−
1ngOrder：ttPPLCa−Cmbro・a77aHs−Simillimaqmandarinia．Thus，thedurationofthereproductiveperiod
lastsaboutamonthineachspecies．
6．Hibernating period
Tbemalesof挽甲8nyabout知ra短wweeksa魚erleavlngtbenest，VisitingFlowers，treeSapandsome
rnushrooms，Whereasthenewqueensleavestraightawayprobablysearchingfbrahibernaculum．Matingoccurs
14
Makoto MÅTSUURA
aroundthenestentrancein Vmandariniaandelsewhereawayfromnestsinotherspecies．Onlyin Vman−
dariniadomalesvisitnestsandawaitdepartingqueensattheentrancetocopulatewiththem（Fig．7）．At
thistimemanymalesof Vmandariniaswarmintheairtoseizethequeens．
Hibernatingqueenssofardiscoveredwerefoundmostlywithinthe50ilorrottenwoodwithahighpercenr
ta＄eOfmoistureasshownin7hble5・Acommonhibernatingsiteforthesespeeiesisinthesoiloverhanglng
Fig．7．Copulatinginlセspa111a（1dori11ia．
Thble5．Hibernationsitesoflセspaspeeies
ぬ
帽
H
Rotten
別
Species
′0
2
0
1
2
247
′h︶
143
8
lO2
7
■．1
Total
l
r．什qP血
Qノ
1▲
y，椚α〃（ね′f〃ぬ
Qノ 0 5
′−U l
4
I∴ロ／柑／′∫
4
O
5
r．αめJ℃
′0
【∴∫血両／／／椚打
′0
7
0ノ
5
勒od
5
Fig．8．t′旨spo（ropIcaqueenhibernatingindecaycd＼＼・00d．
Cl汀飴inthefbrest．But，SOmequeenSOf㌣ざ∼椚′〃∼摺仏巨α〃α胎and HJJ℃ク如wereobservedhibernating
Withirltherottenwoodofoldstumpsorlogslyingontheground（Fig．8）．Theenvironmentsofthesehibeト
nationsitesarealwaysdimandrathermoistplacesintheforesい110Stlyfhcingtothenorth＼Vherethechange
Ofdayandnightternperaturerangeseemstobeleast．Thecellpreparedforhibernation（Orhibernaculum）
andthetunnelleadingtothecellareclearlydugbythequeenherselfandarenotprc−eXistingones．The
insidewallofthecelliscarefu11yfinished；thesoilorwoodsurfacetowhichsheclingsisalwaysmadequite
even．Thetunnelispluggedwithearthorwoodchipsfromwithintocutoffthespacefromtheoutside．
Thehibernatingqueenisal、、′aySfoundclingingtotheupper、、・al10rCeilingofthecell，Closelydrawing
herantennaeandalllegstoherbodysurfaceandconcealingtothetipsofherfoldedwingsundertheventral
Surfaceofherabdomen．Thequeensalwayshibernateonepercell，eXCePtaSlnglecaseof Vana梅where
twoqueens＼＼・erefoundinthesameceLl＼l・ithinrouen＼＼′00d（M八丁SUUR＾1966），thoughS∧Nt）じM＾NN（1936）reported
ComparativebioJogyoftherivcJapanesespeciesofthegcnus LJhpa（Hymenoptera．Vcspidac）
15
thattenqueensofVcrubrvhibernatedtogetherinsidetherottenwoodofadeadashtree．Thesehibernating
habまtsd汀托㌻鉦om物〟ぬ（SÅNDEMÅNN1936，DuNCAN1939，THOMASl粥0）and紬肋錯（MÅTSUURÅ1966）in
theirpredominantlysolitaryhibernationandpreparationofhibernatingchambersinsteadofmereutiliza∼
tionofpre−eXistingcavities．ThequeenswhichenterhibernationhaveovariesofverylmmatureCOndition
andthereisalargeamountoffatbodycoveringtheovarywithintheabdomen（MÅTSUURA1966）．
Thedurationorthedormantperiodis6to8months，VarylngWまththespeciesofhornet；thequeensof
VlrqpLCaSpendtwoorthreemonthslongerinhibernationthananyotherspeciesinsouthernJapan，
Il．NESTING HABITS
ThenestsofhornetsareremarkableandcomplexstructurewhichrequlreagreatprOpOrtionoftimeand
energylnCOnStruCtionandmaintenance，Theearlypartofnestbuildingisperformedbythefbundingqueen，
butthelargerpartofitisaccomplishedbytheworkers．
Inthischapter，aSpeCtSOftheirnestinghabitswhichhavemanyspeciaifbaturesarediscussed，tOgether
With somespecificdifferenees．
l．Nest sites
AllJapanesespeciesof挽汐aaredistributedinmountainoutskirtsandhiilsides，beingrelativelyscarce
plainsandinhighmountains．Nestssitesof掩甲aareShowninT汝ble6．Baseduponthedataobtainedfrom
取ble6．Nestsitesof物αSpeCies．
Coveredpiace
Openpはce
Bough or Ea＼′eS Or Rock Olller
Thicket tree building wa】1 5ites
Ho”0、、・ Olher Under昌一o−
Altic lree Sile5
und ’恥臨1
CaYily
0
24
3
8
34
73
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39
2
3
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178
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164
0
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rt用α〃（ね〟／才ぬ
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0
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ト1∫r〃J〟JJ〃川
2
95
0
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r．α〃α地
39
4l
Notes：A＊，neSt5itesbeぬreorwithouもre王ocationornests；B＊＊，neStSitesar【errelocationornest5．
Fig．9．Nestsof掩軍〟α〝α／お：A，yOungneStinthecooperativeperiod；B，matureneStinthepolyetbicperiod．
16
Makoto M＾TSUUR∧
atotalof596nests，itisrecognizedthatthenestsiteprefヒrenceo‖頻押differscharacteristicallywithspecies，
beingclassifiedintoopenplaceandcoveredplacetypes，Thespecincdifftrencesareasfbllows：Vana飴and
VsimiHimabelongtothespeciesoftheopenplaceprefbrenee．Vana飴nestedonlyinopensituationsabove
groundand1490utOf164nests，Or90．9％，hung伽mbushesortreesneartheground肝ig．9）．Vsknillima
Oftenmovesnestsafterworkeremergence加mtheiroriginalsitestonewlocationsasdiscussedindetaillater．
Thequeensofthisspeciestendtoinitiatetheirnestsincoveredplaces（35／73nests，Or50，7勒）orunderground
CaVities（34／73nests，Or46．6％），Wherethemicroclimateismostsuitablefbrtheearlystageofdevelopment．When
thenestbecornestoolarge，thewholecolonymovestoane叫mOreSPaCioussite，and1320utOf215nests，Or
61・4恥，We代attaChedinopenplacessucha5b払mSOreaⅥ駕Ofbuiはhgs（95／215nests，Or44，2勒）騨ig．10），血ough
itisobviousthatthisspeciesismorevariableinnestsiteselectionthananyothercongenericspeciesinJapan
bothbefbreandaftertherelocationofnests．Ontheotherhand，threespecies，Vc7Vb7q ViTqPicaand Vman一
血rihiqutilizeonlycoveredsituations，VcTUbTVand Vtrppicamaketheirnestseitherinrestrictedcavitiesor
Fig．10．Mature nests o‖句卯SilTliI／i111amade undertheea＼′eS．
ComparativebiologyofthefjveJapanesespeciesofthegenus挽甲a（Hymenoptera，Vespidae）
17
Fig．11．Nestsof Va5PaCTt7b7V；A，neStSiteinho11owtree；B，yOungneStinattic；C，maturene＄tinattic．
inamplespacesbothaboveandunderground（Figs．1l−14），thoughtheformerspeciesoftenmovesitsnest
Site．Vmandarin由nestsonlyundergroundsituations（Fig．15）：amOng56nestsexamined，47werefbund
insubterraneancavitieseitherformedaroundrottenplnerOOtSOrmadebysma11rodents，Snakes，etC．，and
9in subterranean tree hollo＼VS．
Heightofthenestsites abovethegrounddiffcrs、、′ith speCies，lてa／taIisusua11y buildsits nestat10Wer
levelsabout2mfromtheground（14Ooutof164nests，Or85．4％）．Ontheotherhand，1840utOf215nests，
Or85．6％，OftheaerialnestsofVslmillimawerebuilt2∼7maboveground．Nestsofthespeciesgrouppreftrring
COVeredlocations，Vtrppicaand Vc柑b7戦Weremadeatlowcrlevelswithinabout4．5mfromtheground．
Subterraneannestsoffburspecies，Vsimil］imq V trqpicq，T（ctYlbrvandlてmandarin桓Wereal1located
Makoto MATSUURA
柑
Within O．6m be10W theground surface．
Noparticulardirectionalpreferencesfbrnestsofanyofthespeciescouldbedetermined（MATSUURA1971a）．
Fig．12．Maturenest of停5PaCmbro：A．nestin attic；B，bottom ofsame nesl．
ConlParati、■Cbiolog〉・OfthcfivcJapancscspeciesorthegenusレと3Pa（H〉・menOPtCra，Vespidae）
Fig・13・Nestor抱卵αけ叩加：Å，maturemeSt；B，Samel−eStWithenvelopepartiallyremoved．
Fig．14．Nestor掩軍α細〆Cαunde柑rOtlnd．
19
20
MakotoMÅTSUU鼠A
Fig．15．Nestsor挽岬α′77α〃血イ′7ぬunde咽rOund‥A，yOungneSt；B，matureneSt・
ComparalivebiologyortberiveJapanesespeぐiesofthegenus物α（Hymenoptera，Ⅵspidae）
21
2．Nest structure
l）queennest
Concernlngthenestfbundationbyqueenso＝頗卯Onlyasmallamountofdataareavailableonanest
OfVana／ね（A7101）whichwasobservedfromitsveryinitiation（Fig．16，A∼D）．Unfbrtunately，thiscoIony
Fig．16．んnInitia15tageSOfne5tbuildingbya挽卿＝冊曲queen．
WaSOrphanedbythedeathofthequeenbefbretheemergenceofworkers．Fig．17，A・Hshowssomeinci−
pientnestsof打∫／研肋”αWhichiれdicatesuccessivestageshearlynestconstructionbyqueensortbisspecie5．
ThefbllowingdescrlptionisbasedmainlyupontheobservationsglVeninFigs．16and17togetherwithrnany
fragmentaryobservationsin otherspecies．
Thefirststepofnestconstructionbyaqueenistomakeapediceltosupportasmallcomb．Afbw
loadsofwoodpulpareappliedatonespot，fromwhichapedicelismadeintheshapeofaclub，pOinting
Sharplydownward・Assoonasthefirstceu，beingcircularincrosssection，iscompletedonthetipof
22
Makoto MÅTSUURA
A
B
・−、
㊨ H
・
⑳
∴
鰐）
1cm
Fi各．17．Å−H．Initialstagesornestbuildin畠i‡1di的rentnestsor蝮叩α∫如才′J〝〃プ仇
thepedicel，thequeeninscrtsherabdomenintotheceu，andiaysthefirstegg．Whenthefirstcomb，COn−
Sisti喝Of2−4ce重is（usuaily3ceiisin打♂〝鵡and巨J′叩ねα），ha5beeninitiated，血equeenbeginさ拍
COnStruCtanenVelopetocoveritbytheendofthefir＄tday．Thequeennestsof物aspecieshaveusually
3to5cellsaftertwodays，4to7cellsafterfivedays）and6to10cellsaftertendays．Thequeennests
nevercontainmorethanasinglecombandthebasicfぬturesin物αare，incomparisonwithothergenera，
掩5Pu／aand Do／ichove5PLLIa，aS fbllows：
a）托dicei
In掩軍αneStSthepedicelismadecentrallyneverlatera11yasinsome馳ぬ′喝andthecombalwaysassumes
ahorizontalpositionasintheothervesplneWaSpS．Thepedicei，Whichisnrmlyfixedtopreventthenest
frombeingswung，1Sintheshapeofaclub，likethatofPolisteswasps，andstrengthenedbypalnting
alustroussubstanceonitssurface．Ontheotherhand，thequeensofboth物uhiandDo／ichove軍uh7
ぎirstcons汀uciatriangularhangingsheeとOfpaper（ORM￡RO王）1868，DuNCAが1939，丑RIANandBR】AN1948，
SHIDÅ1959）・Fromthecentreoftheloweredgeofthissuspendingsheetathindownwarかpointedspindle
Withaslightspiraltwistisbuilt，andacombofcellsishung什omthetipofthespindle．According亡O
SH】DÅ（1959），’’tbetrian脚はrplate−1ikestalk”whichisusedin掩軍〟由ノ払vね呼ぶuSua11yhasa90∼1800
twistande胎ctivelyabsorbstheimpact裏VentOthenestwhenthecombisswung．
b）Cell
ln Vaspaspecies，the ce11s constructed by queens are always smallerin size than those constructed
ComparativebiologyoftheriveJapanesespeciesofthegenus物α（Hymenoptera，Ⅵspidae）23
職ble7．SizesortlleneStSOr掩堺βSPeCies．
Suspensoria
Species
11∫J〃J／／〟〃J（J
トニ Llr〃わ／・r）
Builder
勒rkercell
Thickness No．ornests
Length Thickness Diarnetcr Depth Thickness ofenvelopc observcd
qしIeen 7．6±0．7＊ 2．3±0．5＊ 7．7±0．2＊ 24．3＝ヒ2義4＊0．18土0．09＊0．34：土0．∂＊ 12
Worker 7．3土0．9 18．6士17．5 乳1土0．9 24．4土2．5 0．28。と0．06 0．48土0．4 10
queen 9．1土0．6 1．9士0．7 8．5土0．3 25．3土1．6 0．24よ0．05 0．23土0．6
Worker 8．8土0．5 10．6土9．7 9．4＝と0．9 28．9土3．2 0．29土0．05 0．34土0．7
仁〟m晶†
queen 7．6去0．4 2．7土0．7 7．3土0，5 23．5土3−10．24土0．07 0．48＝と0．9
Ⅵわrker フ．2土0．6 14．7土9，9 9．8土0．5 25．2土4．6 0．33土0．09 0．58土＝
11／〃〝／l〟（J／・／／血J
queen i5．8土1．】 3．4土0．5 iO，5士1．7 31．6土4．7 0．3i土0．08 0．85：土1．2
Worker 18．9土9．8 5．4土4．8 12．6土2．1 34．2土3．10．40土0．09l．05土0．7
卜二JJ■r）／フ／ぐ（一／
Queen 17．0土1．2 2．3土0．6 7．8よ1．2 23．2土2．3 0．28土0．07 0．68土0．4
Worker 17．6士1．8 6．4土3．4 10．3土1．8 23．6土3．3 0．32土0．05 0．78土1．5
＊：Mean 土 S．d．
1atcrbyworkers，andalsobothceuwallsandenvelopearedistinctlythinnerasshowninrrable7．Since
Ce11sizedirectlya約ctsthebrooddeve蜃opment，thelarvaerearedinsuchsmallcellsemergeasthesmallest
individualsthroughou＝血eseason．NoconsplCuOuSdifferenceintheformsofcellandcombareexhibited
eitherintrかOrinte柑enerica11yamong挽甲q伽〟ぬand加／fcわove堺〟ねatthisstageordevelopment．
ThentImberofccllsbuiltbyasirlglequeenof掩5PaSpeCiesisshownin職ble8．Theseresul【S，a10ng
Withothcrobservationsonmanydesertednests，indicatethatinalltheJapanescspeciesof掩軍a，aSlngle
qtleenCOnStruCtSOnaVerage35∼40ce王1sandrearslO∼151arvacupto【hepupalstagebyherself．No
markeddifftrenceisnoticcdamongthevariousspecies．Ontheotherhand，JÅNET（1903）recordedaqueen
nestof陀5PaC7VbfVCOntaining60ce11sa1loccupiedbybrood，Ofwhich13larvaehadbeenrearedto
pupae・AsfarasIamaware，thisisthemaximumnumberofce11smadebyaslnglequeeninallthe物a
nestsso伽studied．Ⅵvoqueensof掩spao［・ienla［isrearedbyDARCHEN（1964）maderespectively16and
14cel】sunderlaboratoryconditions．In挽軍〟血speciesS111DÅ（1959）descr加danesto‖触血射紬血御
With40cells，andDuNC＾N（1939）rccordedaprobablyincompletenestof掩5Pu血pensyIvanicawith12
andanotherofthesamespeeieswithatleast17．inDolichovespuぬthesameauthorreportedanest，
PrCSunlablyot、DIM（・L（kt［（I，With18cells，andBRuNandBRl∧N（1948and1952）ciledsixnestsorDり・h，e5（／・is
With15，21，31，33，37and42cells，reSPeCtively．Judgingfromtheserecords，thoughstillinsufficientboth
innurnbersofnestsandspeciesobserved，itseemsreasonabletoconcludethatthenumberofcellsfound−
edbyqueensof掩spaspeciesdoesnotmarkedlydeviatefromthatof掩spuktandDo［ichovespu［a．
C）Envelope
Theenvelopeisstartedfromthesubstratum，Orthebaseofthepedicel，neVerinthemiddleofthepedicel．
Specificdiffbrencesintheformofenvelopesaredistinctlyshownamongqueennests．Thefbrmofthe
COmpletedenvelopecanbedividedintothefbnowingthreetypes；bowトshaped，ballqshapedandflask−Shaped．
BowLshLPed功鱒 ThistypehasaninvertedbowトIikeenvelopewhichnevercompletelyenciosesthe
COmb・VITVPica（Fig．19）and V manddrinia（Fig．18）buildthまstype；theenvelopeofthefbrmeris
5．0−5．5cmin diameter and that of thelatter7．0−7．5cm．
Ba〝MShLPed押e：ThistypehasaroughlysphericalenvelopcompletelyencloslnBthecombexceptfbr
nsma11hoIclel、IaLlhebottom．l／：cr〟bro（Fig．20）andト：si）11iI／ima（Figs．17and21）buildncstoft】1e
SametyPe，thoughthenumberofsheetsconstructedvarieswi仁hthespccies．V：c7Vb）・Oa王waysbuildsan
envelopccomposedofasinglesheet，While V：simi！Iima（Fig．17）buildsasecondsandthirdshect，eVen
SOmetimesasixthsheet．Theenvelopeof Vc］VbfVmeaSureS6．8−8，5cmindiameteratitswidestpart，
alld that orト：simiI／iJ7ta4．4−J．6cIllinsidc alld7．5crn oulside．
24
Makoto MATSUVRA
「払ble8．NestcontentsandnumberofcellsinI色甲aSpeCiesattheinclplentStage．
d
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g
cells
H∴扉J〃f／／′J〃α
T． No．of
no．of adults
Larvalinstar
Date Empty
E
Species
F7ask−Shqpedりpe：Thishasacylindricalspoutatthebottomoftheenvelopecomposedofaslngle
Sheet（Fig22）．IntheJapanesespecies，Vana［Lgand VquinLsbuildthistype；theformofthespherical
partoftheenvelopeisslightlyvariable．Of42incipientnestsof VanalLgco11ectedfrom1964to1975，
thediameteratthesphericalpartwas7．4cminthelargestnestand6．lcminthesmallestone（OnaVerage
6．6cm土S．d．0．6cm）．Thecylindricalvestibuleisbuiltbefbrethefirstlarvabeginstospinitscocoonand
isnotrebuiltifdestroyed．Inbothball−Shapedandflask−Shapedtypes（Figs．16and22）thenestsinthe
courseofconstructionvarygraduallylntheshapeoftheenvelopefromaplate−Shapedenvelopeatthe
beginnlngtOthefinalfbrmdescribedabove，becausequeenstendtobuildcellsandenvelopemoreor
ComparativebiologyortlleriveJapanese5peCiesorthegenus挽甲α（Hymenoptera，鳩spidae）
Fig・lS．lniliaLslagCSOrnCSt bし】ildingb〉′nIセspa111L7／Tdori17iL7queCn．
Fig．19．Quee】1neStSO＝句卵彷彿加
ヱ5
2（§
Makoto MA′rSUURA
Fig．20．Queennestor物αC撒か敬
Fig．21．queenne鮎d‖句叩正面肋閥
ComparativebiologyortheriveJapanesespeciesorthegenus掩軍α（Hymenoptera，鳩spidae）
27
Fig，22．Nestsof物qanu／is：A，B；COmPletequeennestswithatubularvestibu王e；C，neStaftertheworkers
ha＼でrCmO＼′edlhelubular veslibule．
lesssimultaIleOuSlyuntilreachingthe finalslagC．
2）Changeinnestshape雨ththeemergenceorworkers
Afterthefirstworkersemerge，theshapeoftheenvelopechangesandquicklylosesthecharacteristicnest
ShapelIn VshniHimawhentheworkersbeginaddingnewscale−Hkelayersontheoutside，theyalsotearaway
theinnerlayersmadebythequeentomakece11sfbrtheenlargementofthecomb．TheHrstfbwlayersinside
thenestarechewedawayuntilonlyadiscrlikeremnantremainsabovethecomb，irlCOntraSttODoHchovがpu由
nests，Wheretheenveloperemainscomposedcssentiallyofseparatelayersofpaper（DuNCAN1939）．ln Vana船
theworkersmakenoadditionstothecylindricalvestibuleatthebottomoftheenvelopebuiltbythcqueens．
A鮎rfburornveworkershaveemerged，thistubularpassagewayiscompletelyremovedbytheworkcrsandisnot
Makoto MATSUURA
2る
reconstructed（Fig．22．C）．Theenvelopeisenlargedbytheadditionofnewscalesontheoutsidebyworkers
andtheentranceisshiftedfromaventraipositiontoapositioninthedirectionwhichaffordsthecasiest
accessto foragingworkers，Ontheotherhand，theworkersof V Erqpicaand V mar7dariniabuild an
additionalwalllayerdowntothefirstenvelopeeonstructedbythequeen，andremnantsofearlyenvelopes
remainabovethetopmostcombwithoutanydisturbancebyworkers．Vc7t7broworkerstakeofftheoriginal
envelopeexceptforaplate−Shapedpar＝eftatthebaseofattachment（Fig．23），andthisfbrmismain
Fig，23．1blユngne5tOr掩卿α仁和かα
【ained untj】【heend ofJul）・．Such treatrncnt seemsto beaspeciriccharacter ofthisspeeies．
3）Developed nests
S＾USSURE（1853−1855）dividedthenestsofsocialwaspsintotwoprimarytypes，gymnOdomousandstelocyト
tarous．Vtspinenestsbelongtypicallytothelattertypewhichisenclosedinanenvelopeandcontains
SeVeralcombssuspendedonebelowtheotherand＼Vithoutconnection，＼＼・iththeenvelopc．Malurenests
Of掩軍aeXhibitcertainspecificdifferencesinneststructure．Basicfeaturesaredescribedbelow．
a）Pi11ar
lnlセ5PaneStSeXCePtforレ：alta／iseacheombisconnectedtotheothersbyonemai11pillar（0．8−2．OcI11
long）and2∼50auxiliarypillars．Inthenestof Vana掠thefirstcombalwayshangsfromthesupport
byonepedicelandadditionalcombsarealsohungfromtheearlieronebyapedicel．Asnestsofa11掩5Pa
SpeCiesareenlargedandtheir≠・eightincreases，additionstothepedicels aremade b〉T Plasteringpaper
pulponthemandontheadjacentsurfhcesofthecombs．Thistreatmentthickensthepedicelsandbroadens
theirbases．lnthecaseofIてsiITti／IiI77athesizeofpedicelsbuiltabo＼・ethcrirstcombreaches5−9cmin
thickness．
b）Cells
Theaヽ■erageSize ofbrood eellsin 停spaspcciesisgi＼・・eni111bble7．Thesize of＼＼′Orkercellsis ranked
asmL7／Zddri／）iG−［lPPica−CIVbro−L7／1a／is−5imj／／i／71L7，thoughthecellsfor＼VOrkersandsexualsaredifferent
evenineachspecies．fnailJapanesespeciesceilsbuiltbythequeenareused2∼3tirnes；uSageOnlyonce
OrmOrethanthreetimesisso farunknown．Incombsbuiltbyworkers，CeuSOftheuppercombsare
used2∼3timesandoftheloweronesl∼2times．Buteveninthetipcombsomcperipheralincomplcte
Cellsappeartobeunused，judgingfromtheabsenceofcocoonresidues．
ComparativebiologyofthefiveJapanesespeciesofthegenus掩箪a（Hymenoptera，Vespidae）
29
C）Conlbs
Inthreespecies，Vma71dariniq Va77aZi5and Vtrqpicqthedorsalsur蝕ceofeachcornbisconicaland
higheratthecentre，Thesurfaceisroughandthebottomofeachcelliseasilyrecognized，COntraSting
tothehorizontalcombswith nat，eVendorsalsurfhcesinIイcrabroandlてsimi［［ima．1nIてmandari／7ia
thesynchronousbuildingofafewprimordiaandtheirlaterfusionformingasinglecombisnotrare，
butintheotherfourspeciesanewcombstartsalwaysfromaslnglecentralprirnordiumwhichisgraduaト
1〉′eXpandedeoncentricallyiFnotobstructed．
Thenumberofcombspernestatalateactiveperiodis4∼7in‡てmandariniち7∼10in V
6−7inIてcrabrq3−4inlてana／isand2−3in V（rOPica．ThetopcombofIてmandar［niaandトて（rOPica
isabandonedaたersummeちa魚erwhichitisattackedbymoulds，Whereas Vc7t7bTVandespecia11y V
and Vsimi〟imastrengthenthetipcombbyfillingemptybroodcellswithexogenousmaterialsandutilize
itasastrongsupportforthelowercombs．Thelargestcombssofarmeasuredwere6l．Ox48．Ocmwith
l，191cellsinIて171a（tdari／血（S6403），39．0×38．5cmwithl，177ce11sinlてsimi／／ima（S6403）．40．5×40．Ocm
、、′ithl，468ce11sinレ：亡用bTD（C7307），16×16cm≠・ith250ce11sinレ：a／沌旭（A7522），and14．0×11．5cm＼＼・・ith
155ce11sin VtJ℃pica（T7105），reSpeCtively．
d）Enve10Pe
Thegeneralshapeoftheenvelopechangesbet＼l・eenthespecies oftheopen placepreftrenceand thoseof
thecoveredplaceprefbrenceasstatedearlier．Inthefirstgroupwith VsimiHimaand Vana貼theenvelope
isusuallythick，COVeringthenestcompletely．Indevelopednestsofthesespeciestheenvelopesconsistofrnulti−
layeredsheu−1ikesheetsinvol＼′ingman），aerialchambersbetweenthem．InthecaseofIイsimi［［imathethickness
Ofenvelopeinalargenestreached3．5∼5．5crnatthemiddleand12cmatthebottom．Ontheotherhand，
the＄eCOndgroupwithIてcTUbrQ，VtTWtCaand Vmandariniatheenvelopeconsi＄tSOfonlyafbwplate・like
Sheetswithimperfbctaerialchambers，andthelowestcombisalwaysexposedwithinthenestcavity．
e）Nestentrance
Normauyallthenestsof挽甲ahaveonlyoneentrancehole．Thenestsof Vsimil［imaand VanaHshave
asinglecircularopenlngShiftinglnPOSitionfromthebottomofthenesttothesideataveryearlystage；
usuallytheentranceinoldnestsisrelativelyhighuponthenestwall．Ontheotherhand，thespecies
bui）dingincovered places，SuChasレ：mandarinia，H c（tlbro（Fig．12）and tイtrqpica（Fig．24），aLways
Fig．24．NestentranceorI々5pαJ′叩fぐ乱
30
Makoto MÅTSUU見入
Omittheenvelopeatthebottomofthenestandtheworkersutilizetheopeningasanentrancehold．In
Subterraneannes【Sthepre・eXistjngburrol＼・COnneCting thecavit），andgroundsurfhcejs used as theen−
trancecanalwiththelength＼・arying from2to60cm．
In VcTVb7VCreVicesaroundthenestaren11iystufftdwithwoodpulp，Whichattainedinahighlydeveloped
CaSe5minlengthand20cminwidth．Suchtre8tmentSeemStObeaspecificcharacterofVcTVbrqthough
【hell・OrkersofIイJrqPicaandlて／T）G／JdGri／7joalsobui】dafe＼Vrudimentar）・Sea】5aItheentranee5tOI】eStS．
3．ModiFication ofnest site
Aftertheemergenceofworkersanestsiterequircsaccrtain amount oflnOdiricationtoadaptit tothe
needsofagrowlngCOlony．Vmanda7・iniaqueenswhichpreferrathernarrowcavitiesunderground，doa
greatdealofexcavatlngtOenlargethecavityinwhichthenestisbuilt．Nestcavitiesareextendedassoil
adjacenttothenestenvelopeisconstantlyremoveda一文dtakenawayfromthenests．Theworkersremovesoil
Particlesandmakeasma11baliwiththeirmandibles．Keeplngtheballbetweenthemandibles，theyleave
thenestonfootanddroptheba11closetotheentrance．Atthepeakofnestexpansionatongue−1ikeplatform
Ofremovedsoilappearsattheentrance，Which，inahighiydevelopedcase，attained2minlength，50cmin
widthand15cminmaximumdepth．Stonestoolargetobernovedaredroppedtothebottomofthenest
CaVitybytheremovaloftheearthbeneaththem．Onthecontrary，in V（7叩icq，Whichbuildsthesmallest
nestamongJapanesehornets，arelativelywidespaceinthecoveredplaceischosenastheirnestsite，SOthat
thereisalwaysaspacebetweenthewallofthenestandthatofthecavity．7（ana偽doesveryiitt￡ework
uponthenestsitebecausethequeennestsonlylnOpenSituationsaboveground，thoughtheworkersclear
a＼、′ayleavesandt＼＼′igsaroundthenestwhenthenestisbuiltin densc foliage．Ontheotherhand，bothIrて
Simi〟imaand‡くごTVbTqWhennestinglnnarrOWCaVities，uSua11yabandontheorlglnalnestsitesandrelocate
thecoIonieswithoutgnawingthewausofthecavitiestoenlargethem．Adetaiiedaccountonthisphenomenon
is gi、′enin the follo＼＼・ingsection．
ヰ．Relocation of nest
Ass（atedin（heprevjoussec（ionll：simiI／iJ71L7be10ngStOthe5PeCiesoftheopenplacepref七rence．AsLO
thequeerlneSt，however；itisclearthatthequeenprefersaconcealedsite（7hble6），thoughismorevariable
innestsiteselectionthananyothercongenericspeciesinJapan．Inthisspecies30outof50nests（60．0勒）
builtbythequeenwererelocatedtoadifftrentlocalitywhenthenestshadattainedlargesizeandrequired
morespaceforcombbuildirlg．AmongthenestsofVsimillimaobservedafterlateJulyfrorn1964till1975，
Thble9．Dateorrirst observation orrclocalingcolonicsorl；＆spusimi／／ilTlaand LてぐrablV．
1托ar
’65
′66
′67
り0
′71
ワ2
′73
JuL．20 Jul．26 Aug．8 Jul．17 Ju）．15
l1∫J〃扉仙朋
・ニ
′68
り5
Aug．2 Jul．ユ2
（S6701）（S6801）（S7003）（S7106）（S7201）（S7304）（S7503）
−
‥
ー
1320utOf149nests（88．6勒）weresuchrelocatednest，In Vc7tlbrqwhosequeenbuildsonlyinconcealed
Situations，24outof45nests（53．3甲b）observedafterJulyfrom1966till1975werealsore10Catedfromthe
Or唱Inalnestingsite．Ontheotherhand，王havenotobservedsuchrelocationofnestsintheotherthreespecies，
Vanaljs，Vttppicaand Vmandarinia・TherelocatednestswereobservedinmidpJulytoearlyAugustin
VsimillimaandinearlyJulytolateAugustin VcTY7b7VaSShowninrrables9andlO・Thenestswithrnany
broodsuggestthatrelocationhadbegunearlierthantheobservedtimes・Judgingfrom7ables9andlO）and
otherobservationsonmanynestssofarobserved，itseemsthatthequeensleavetheirorlglnainestfrom
lateJunein both species．
ComparativebiologyortbenYeJapanese5peCiesorthegenus物α（Hymenoptera，Ⅵspidae）
31
7bble10．Coionycornpositionandrelateddatafororlglnalandrelocatednestsof物asimi［／kTla．
Nest code
Nestil鳩Site
1∴−＿…⊥∴−
combsCe11s
（m）
0＊S7105 Undergrollndcavity
一こ
eggSl＄て 2nd 3rd 41！− 5th 蛍石 ♀ 受苗
Jul・17り13 33135・0 0
32
R…S7106 At【ics
￣
−
．
い・
Jul−Ⅰ7り1 3 368 0．5 147
一−
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Au8，17り2 2 23ヰ 72．6 11
3 2 0 0 】6 37 0 0 ほ（0） 0
R・S7214 Ⅰ≡avesorbuilding
Aug．17りヱ 3 400 0．8 】86
0▲S7302 Under即Oumdeavily
Jul・25り3 2
凸7
Aug．7り3 3
1
0・S7214 Cavityunderrooト
2
︵U
0
0
0
へU
21
17
13
Qノ
00
つ一
つー
8
nV
っ川
8
39
乞
つー
0・S740ユ Under即肌ndcavity
R・S7401 Eavesorbtlilding
lZ
▲l
Eave50rbuildii鳩
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R・S7307
6
′り
in法titles
88
0
49
0 101（8）0
46 72 0 0 20（0） 0
66 154 0 1 22（0） 0
恥】6り4 3 94157・0 36 0 2 515158i87 0 0190（8）O
4
Sep．16り4 5 2，115 77．0 446 46 85 23
2 1 0
28 1140（0）232
．0：Origina）ncst． ◆●R：Relocated11eSし雪二：Foundressquccn．Figuresin parcn【hcsesdcnoleStylopiヱed＼＼・Orkers．
Basedupontherecordsofabout150nestsofboth tJ：silTti／IimaandIイcrT7b］’00bservedfrom1965to1975，
thcprocessofrelocationofthenestisi11ustratedschematieall〉′inFig．25．ubrkersbegintosearchforne＼＼′
Sitewhenthespacebecomestoonarrowfbrtheirnest．Scoutworkersflyinalldirectionsinsearchofanew
permanentnestsite．AsalreadypointedouttheworkersofVsiml〃imaselectaerialsituationsinmostcases，
Whereasthoseof VcrYZb7Valwayssearchforconcealedsituationsfortheirnewnestsite．Whenasuitable
Siteis fbundnot farfromtheor唱InalnestthescoutsettleonthepIaceandspendmostoftheirtimeat
restthere（Figs．26．Aand27．A）．Theyonlyoccasiona11yreturntotheoriginalnest．Different5COutSSettle
Ondifferentsitessimultaneously，andacontestensues．Selectionofanestsiteiscontinuedforseveraldays
andatthemostsuitablesitethenumberofworkersincreasesduringtheperiod．Withthecommencement
①
● ＝FOい」D川G Q〕∈EN
O 王 OLD 吊ORKERS
O ；YOUN6 日ORKERS
∧ −‥・・‥−・、
E：EGGS
AND MALES
」‡LARVAE
●
●●●●●
000 00
0息
E し P
P！PUPAE
Fig．25．Thcproccssorrelocalio】1OrCOlonicsi】1卜：si／1）i／／ima an〔1tlcrabl■O1
1．Ne芳一払undationbyqtleeninnarrowcavity．2．Withthegrowthort‡1eCOiony，thenestrequlreSmOre
a＼・ailablcspaee．3．Scou（＼＼・Orkersbegin toscarch roraIle、＼・neSISilCnearlhcoriginalncsl．4．Whcn
asuitablesiteisfoundthescotltSSettleondifferentsitesandacontcstensues．5．Selectinganesting
Sitecontinuesforseveraldays．6．Thenumberofworkersincreasesatthemostsuitablesite．7．Thc
Oldworkersandthequecnleaveindependentlyto5earChforthenewnestsite．8．Aboutadayafter
arri、′alorthcquecIlthe“・Orkcrsslarlbuildingane、、・eO【11bandthequcenla）・SeggS・9．Mos10rthc
WOrkersinthenewnestrrequentlyvisittheoldnestto托edtherelnaini‡lglarvaeandyoungworkers
untiltheyleave．10．Newlyemergedworkerssearchfbrtheirnewnest．11．Theoriginalne5tisaba‡1dorト
ed when no brood remain to be托d．
32
Makoto M＾TSUUR＾
Fig．26．Nestrelocationin fespasbllH［ima：A，WOrkersandthcqueengatheringatanewsite；B，COnStruC−
tionornew cells．
Comparati＼・ebiolog）・O［thcriveJapanescspeciesorlhegenuslセ5PO（H〉・menOPtera．Vcspidae）
Fig．26・Nestrelocatio狛血掩即射由痛仙川 C−neStW氾＝＝udimen【aryenVelope；DナneStWithaspherical
cnvelopcenc10Singthcrirsteombl
33
34
Makoto MATSUU鼠Å
Fig．27．Nestrelocationin噸ac7Vb7V：A，WOrkersgatheringatanewsite；B，COmpletednest．
Ofsearchingforanewnestsite，WOrkersintheorlglnalnestceasebuildingactivitiesandthequeenstops
OVlpOSiting・ThensheleavesonherowntosearchforthenewrleStSite・Aboutadayaftcrarrivalofthequeen
theworkersstartbuildinganewnest，Assoonasthefirstcombconsistingoffivetotwelvecells，Without
Comparati＼でbiologyorthcrivcJar）aneSCSPCCiesofthegcIluS咋spL7（Hymcnoptcra，＼七spidae）
35
anenve10Pe，isconstructedthequeenbeginstolayaneggineachcell（Fig．25．B）．Withinseveraldaysworkers
COmmenCetObuildaroughlysphericalenvelopecomposedorasinglesbeet（Fig．25．C）andcoverthecomb
COmPletelyexceptfbrasma11holeatthebottom（Fig．25．D）．Thesecondcombisbuiltafterthefirstone
iscompletelycoveredwithenvelope．Thequeenstaysatthenewnestandneverengagesinextranidalactivities．
NowtherearetwocoloniesusuallylOmorlessapart（althoughthemaximumrecordedwas180m）．The
fractionremainlngattheorlglnalnestcontainsyoungworkersandbrood，andthefraetionatthenewnest
COntainsolderworkersandthequeenwithoutlarvaeandpupae・Inadditiontotheirtasksinthenewnest，
mostoftheworkersfrequentlyvisittheoldnesttofeedlarvaeandnewlyemergedworkersuntiltheyleave
indcpendentlyforthenewnest．Thus，theworkersemerglngintheoldnestperformorientationflightswhen
twoorthreedaysold（probablyassoonastheyareendowedwithflyingactivity）andthenstarttosearch
fbrtherelocatednest．ThusrelocationusuallylastsforaboutamonthuntilthemaJOrityofbroodmature
andmovefromtheoldnesttothenewone．Inthecaseof V：simil〟mathedateofthelastobservationof
relocatingworkerswasAugust9thin1971（NestS7109）andAugust29thin1968（NestS6704）．Withtheemergence
Ofworkersinthenewnesttheorlglnaloneisabandonedbecausethereisnolongeranybroodtofted．
Thedescriptionsgivenabovesuggestsomebionomiccharacter5SPeCificto巨∫′椚∼J伽7αand巨c和∂和Which
CauSethemtovacateoldsitesandconstructnewnestselsewhere．thisbehaviourmaybecausedbythelimited
SpaCeintheorlglnalnestingsite．Asalreadystated，thequeensofthesespeciespreferrathernarrowcavities
atthetimeofnestfoundation（Thble6）andtheworkersscarcelyextendthecavities．Incontrast，Vman−
darinjaworkersconstantlyremovethesoiladjacenttothenestanddeposititawayfrornthenest．Thereason
thatthequeensofthesetwospeciesselectsuchnarrowspacesfbrtheirprlmaryneStSitesseemstoberelated
totheirrelativelyearlyappearanceandnestfbundation（7hbles2and3）：themicroclimateinthesoilorin
thesmallcavitiesappearstobeclosetoidealforbrooddevelopmentintheearlynestingstage．Furthermore，
COlonyemlgrationinthesespeciesisnotparticularlyprecariousundertakingbecausethebroodleftinthe
Oldnestarerearedsuccessfullybyworkerslocatedtothenewnest．Withtherelocationofthenest，theco10ny
Canentertheexpansionphaseofitscycleinthenewsitewithmorespace．
Ontheotherhand，SOmeSOCial＼VaSpSandbeesarealsoableto＼′aCateOld，unfavourablesitesandcon−
StruCtneWneStSelsewhere，Whichiscalled“absconding”．Thebroodcannotbemovedinthiscase，andit
isleftbehindtodie．Inthemeliponinebees，abscondingisarareandespeciallyprecariousundertaking．When
itoccurs，aSrepOrtedin7埴OnabyPoRTUGALrARAUJO（1963），forexample，thequeenmustbeieftbehind
Sincesheisunabletorl某andtileCOIonymustproduceanewqueenandseehersuccess餌Iythroughher
nuptialflightinordertosurvive．
5．Thermoregulation
Theregulationornesttemperatureisageneralcapabilityorthesocialinsects．In掩軍αCOlonies，When
WOrkersareabsentorfewduringtheearlypartoftheseason，thequeensandworkersrarelyperformtheト
moregulationandthetemperatureofthenestapproximatesthatofthesurroundings（rrhblell）．Withthe
developmentofthecoIonyandthebuildingofthickerenvelopes，thepresenceandtheactivityofthenumerous
adultsandlarvaeinthenestcausesacertainelevationofthenesttemperatureascomparedtothatofthe
SurrOundingenvironment．Fromearlysummertoautumn，COIoniesareabletoregulatetheirtemperature
asshownin一誌blellandFig．28．InthecoIoniesof掩軍aSpeCiesthetemperaturewithinthecolonyusuauy
Stayedcloseto320Cduringmostofthesummerdaysandca．3OOCduringearlyautumn．Thediurnalrange
OfnesttemperatureonadayinearlySeptemberwas29．9∼30．60Cin掩spasimil［imaand28．5∼30．20Cin
Vana［isasiuustratedinFig．29．AccordingtoIsHAYetal（1967）anestof Vorienta船inthegroundwith
atemperatureof28．50CnuctuatedlessthanonedegreecentigradedespiteanexternalvariationoflO∼390C．
HIMMヱR（1932）statedthattheinternalnestternperatureof物uhiVuなarisremainedalmostalwaysbetween
29．50Cand32OCthoughtheoutsidetemperatureaveraged18．5OCduringtheobservationperiod．Thesedata
indicatethattemperaturecontrolinwaspcolorliesislessconstantanduniformthanthe35OCofahoneybee
COlonywhichfluctuateslessthanO．50Cduringtheactiveseason（GATES1914，HIMMER1932，LIND＾UER1954）．
The regulation of nest temperatureis attainedin severalways．The elevation ofnest temperatureis
Makoto MÅTSUURA
36
1もblell．Nesttemperaturemaintainedbysomecoloniesof掩5PaSPeCies．
1b】lP．（OC）or
Species
Date
Nest
20．重
Qノ
︵U
27．4
21
二・
31
3
5
1
30
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30
−￣
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l繁，3尊
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1
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22
24
4
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け
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LL﹁ん
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16
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25．5 12O壬
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Undcrground
1♀
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24．2 1翠，8登
28．3 90苧，4谷
Thicke1
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Thicket
22．3 1学，60肇，11♀ Thicket
王6．4 39萱，36苔
lO．3 33馨
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7
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No＼・．ヰ’7l
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7
7
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Aug．20’73
Sep．24’68
′0
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3
7
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May 29’68
Jul． 8〉68
︵U
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Nov．25，67
3
2
Jul．1了’7i1
0ct．3lI67
H′J叩ね〃
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29
Jurl．29’75
三aves
23．0 1学，875蛍 Underground
8
MaylOり3
Inside bee hive
1単
26．2 1♀，92単
8
J山． 9，71
．亀ug．19り1
Sep．15’71
0ct． 5■70
ト：J〃α／！d打「血血
No．or adulls NesISite
2
May 26り3
レ：∫J椚〟〃川α
Out5ide
23．0
1翠
1竿，10登
27．5
30．1 1ギ，15‡
Underground
Underground
underground
Thetemperaturereadingsweretakeninthecentreofthenestandintheshadeorinthecavityinthevicinity
Or e8Ch nest．
probablyattributabletothenaturaloutputofheatwhichisgeneratedasaby−prOductofmetabolisminthe
colony．AccordingtoCHÅUVIN（1968）ahoneybeegeneratesatleastO．1calorieperminuteat10OCundermost
conditions．In挽野aCOlonies，tOO，aCertainamountofheatwouldpresumabiybeproducedbyallthemembers
ofthecolonies．IsHAYandRuTTNER（1971）reportedin rc用わ和COloniesthatthemovementso‖arvaeand
theirmetabolicaetivitiescouldcreatesufficientheattomaintainreasonablethermoregulationinnestseven
intheabsenceofadults，andinpupaleellsaconstanttemperatureofca．30OCwasmaintained，although
diurnalfluc（uationsinlar＼・alcells＼VaS muCh greater．
Whenthenesttemperaturereachessomepointaround300CtheworkersbegintofhnwiththeirwlngS
insidethenest．Ifthetemperaturecontinuestorise，mOreWOrkersengage主nsuchalarventilationnotonly
insidethenestbutonthesurfaceoftheenvelope，andsomeworkersbegintobringwaterwhichisthendeposited
asdropletsonthecapsofcocoonsorthepupalcells．Otberworkerssuckthedropletsorwaterandthen
spreaditintothinfilmsonthecornbsurface，eXtendingthetonguesoutward・Asotherworkersfanwith
theirwlngStOdrivemoistairawayandoutofthenest＞thewaterevaporatesandcooIstheairinthenest・
Similarbehaviourofbothfhnnlngandwaterdropletstocooloverheatednestsarereportedarnongthecoト
oniesが劫／如錯（STEINER1930），撤少抽（ScHWÅRヱ1931），掩軍〟由（WEYRAUCH1936，OAUL1952，KEMpERand
Dt）HRTNG1967）andthehoneybee（P＾RK1925，LINI）＾UEitt1961）l
Inadditiontothewaysmentionedabove，theregulationofthenesttemperatureisefficientlyaccomplished
ComparativebiologyofthefiveJapanesespeciesofthegenus掩SPa（Hymenoptera，Vespidae）
37
℃
巴⊃−望むd∈ む卜
OCTOBER
SEPTEMBER
NOVEMBER
Fig．28．Fluctuationsofthetemperaturewithinthenestsof物aanalisand Vsimi／［imacomparedwiththe
airoutsidethenestat22：30duringautumn，1969．Thenurnberofoccupantswasca．65Oworkersin
l・て∫imi／／imqcoIonyandca．70≠rOrkersin tイona／i5COIony．respec【ivel〉・，OnSeptember20th．A：neSt
temperattlreOfVsimi（（jma；B：neSttemperatureOf Var2alis；C：airtemperatureinthevesplary；D：
airtemperatureoutsidethevesplary．
0
3
むJ⊃︸のしむnこLむト
／−′
＿−／
ニ
ク∴＼、
′
・
こ、
l、‥＼ごこ−−・−＝ t−
●−（9
6 8 柑12†ヰ】618 20 22 2ヰ 2 4 6T‡ME
Septembe「7
Sep．8
Fig．29．Diurnalnuctuationsofnesttemperaturein掩spasわ7jllimaand V：ana［iscolonies．Thenumberof
OCeupantSWaSea．550in レ：simi／／imaand ca．60in レ：ana／is，reSPeCtiヽ・ely．Notationasin Fig．28．
byaddingnumerousairspacesintheenvelope，SOrnetimesasmuchaslOcrnthickin Vsimi〃imちWhich
helptoinsulatethecomb丘omtheinfluenceofexternaltemperature，In Vorienta梅IsM＾Yetal（1971）observed
3S
Makoto MATSUURA
thatthehornetswallo汀mostorthenestentranceandconstruct＝speciaienvelopesHaroundtbepopulated
COmbswhenthenesttemperaturedropsbe10Wtheoptlmalpointattheendoftheseason：Ihavenotsofar
ObservedsuchbehaviourintheJapanese掩印a・Thenesttemperature，howeveちdecreasesa10ngWiththe
dropmairtemperatureoutsidebecausedeathofworkersaswellasdepartureofthesexualsbringsabout
adiminutionofthepopulationsizeincoloniesasautumnisfaradvancedasshowninFig．28．Asudden
droplnneSttemperatureOfbothspecies，Vsimi］limaand Vana［islattheendofOctoberseemstohave
beencausedbycessationofactivitywithinthenests．
6．Nestdefenceand aggressiveness
All掩spaspeciesaregenerallyregardedasextremelyaggressiveanddangerous，butthequeensofallthe
SpeCiesinJapanarelessaggressivethanworkers．Usuallyshedoesnotattackmanandanimalsevenifthe
nestisdisturbedbythem，thoughsheshowsgreatalarmandrushesaboutontheenvelopeandcombor
fliesaroundthenest・Ontheotherhand，attheinvasionbyalienqueens，鉦equentlyobservedin㌣ご用み和
beねreanda鮎remergenceofworkers（JANETl卯3），allinhabitants，bothqueenandworkers，ruSbatthe
intruderandattackwithmandiblesandstingstokillit（MATSUURÅ1970）．
Theworkersofvariousspeciesshowmarkedlydiffbrentreactionswithregardtodisturbanceoftheirnests．
Therelativeag即eSSivenessofvariousspeciestomenandanimaisatorneartheir錮′nneStSisintheorder：
mandarinia−Sim〟〟ma−CTqbro−ana胎−trqPica・Thefirstthreespeciesaresoonprovokedtoattackwith
regardtoanapproachtotheirnest，aSis VvelutinainFbrmosa．（MATSUuRA1973）．Inmyexperiencethese
WOrkersusua11yattackedandfbllowedthemenwhoapproachedthenestswithinabout5∼10m．Assoon
asaguardattheentranceisalertedbythesightofalargeobjectmovlngnearby，Shemaydartsuddenly
fromtheentrancetothesurfaceenvelope．Thiscausesageneralriseinactivltyandotherworkersinthe
nestbegintodartandfliptheirwlngSOVertheenvelope．Thenestnowbristleswithaggressivehornetsand
anaudiblevibrationthroughouttheentirecolonyiscaused・Ontheotherhand，Vanalisisnotaveryag−
gressivehornetaspointedoutbyVECHT（1957）．Usuallytheworkersofthisspeciesdonotattackunlessthe
nestisdisturbedandonecanapproachthenesttoaboutlmwithoutanyattack．However，Whenthenest
isdisturbedmanyworkershurryoutoftheentranceandruninaudirectionsovertheenvelope．Someof
themnyaroundthenesttocarryoutanattack．巨′J叩∼C〝istheleast喝greSSiveortbeJapanesehornets：
althoughIobservedabout40neststheworkersofthisspeciesdidnotattackmeevenatthetimeofcollecting
thenestwithoutanyanaesthetization，thoughtheyviolentlyflewaroundmeandthreatenedmebyapar−
ticularloudclickingproducedbyrapidopeningandcloslngOfthemandibles．Thisclickingisproducedin
a11物aspeciesatorneartheirnestsandisregardedascharacteristicoforprevalentinthegenus．In V
椚α〃（ねr′〃ぬitisproducednotonlyattbe11eStbutalsoattheoccupiedhivesandnestsorothersocialwasps
andtreesapsourcesfrequentlyvisited．Thisindicatesitwendevelopedterritorialde飴nce．Ontheotherhand，
theotherfourspecies maketheclickingbehaviouronlyatorneartheirnests．
Duringthedaytimeina11Japanese物aspeciestherearenohornetsontheenvelopeexceptfbrtheworkers
engagedinmakingtheenvelopeandin触一nlng・In VsimiHimaseveraltensofworkersaiwaysstayovernight
Ontheenvelopecenteringaroundtheentrance，unliketheotherfburspeciesinwhichtheworkersareall
insidethenestat
aroundthenestwithoutanyflightassoonasanestisdisturbedintheevenlngandnight．Intheotherspecies
thereareseveralguardspositionedatthencstentranceatnight．Theyhurryoutoftheentrancetocarry
Out an attaCk whenIllan Oranimals approach toocloscly．
、ItseemsthatthedefbnsivepowerofacoIonyof掩軍aisdirectlyrelatedtothesizeofthepopulation，
Vanajisand Vtrqpicabelongtothesmallpopulationtype，neVereXCeedingl，000incellnumber，being
equalorsmallerincolonysIZeincomparisonwith撤触gぶSpeCiesoftheIargestcolony，SuChas月ぐ紬e〃め
Psne他niandRrDthnqyi（MATSUURA1970），Ontheotherhand，Vmanda7・iniq，Vsimillimaand Vc7Vbrq
regardedasaggressivespecies，aretObeclassifiedintheiargepopulationtypeandexceed4，000inmaximum
Ce11numberassho、、・nin chap．V．Population dynamics．
Dis（！uSSion
物aextendwellintothesubtroplCSandtropICS，Wherepredationbyantsonnestcontentsisaserious
Compara“vebiologyortheriveJapanesespecie50rthegenus物α（Hymenoptera，Ⅵspidae）
39
threatduringthesolitarynestingperiod，eSPeCiallywhenthequeenisonafbraglngtrip，aSStreSSedbyJEÅNNE
（1975）．Thequeennestsofthcflask−Shapedtypewithacylindricalvestibule・builtbyonlytwig−neStingspecies
OfIてanalisarld Vqmn也essentiauysubtropica王ortropicalspecies，COuldserveasaprotectionagainstpredators
assuggestedbyYAMÅNEandMAKINO（1977）．Asimilarstructurehasbeenreportedinthequeennestsofsome
Do／ichoIJeSPu／a5peCies，SuCllaSDITtaCEl／0（a（R＾U1929，DuNC∧N1939，T＾K＾M】Z∧VrI＾per．COm．）＼＼血ichbuild
essentia11yaerialnests去uspendedfromtreebranches．Astheantsareaseriouscontributortocolonyfailure
inqueennestsofsometreenestingjb／LFteSinthetemperatureregions，tOO（YosHIKAWA1957，MIYANO1980），
thelongvestibuleisregardedasanadaptationforbrooddefenceinvespIneneStSOftreenestingspecies．
Thenestsofthebowl−Shapedtypearefoundinthespecieswitharathersoutherndistribution，SuChas
V：manda7血jdand Vt／叩ICa，Theybuildnestsexclusivelyinconcealedorundergroundsites，Wherethisnesting
habitmayreducethedangerfromantpredation，andassistwiththermoregulationduringthesolitarystage．
TheincompletecoverlngOfthecombsinthesespeciescouldberegardedasadaptationstosuchnidifications．
Ontheotherhand，thenestsoftheball．shapedtypeareonlybuiltbytyplCallytemperatespecies，SuCh
as V：c7VbrDand Vsimillimawitharathernortherndistribution（VECHT1959）．AccordingtoDuNCÅN（1939），
V：crvbrDbuildsanenvelopeifthenestismadeintheopenbutnotifmadeinhollowtrees，eXCeptPerhaps
afewr11dimentsabovethefirstcomb，andsubterraneannestsusuallyhaveenvelopesexceptforthosebuilt
inveryhardandstonygroundinwhichthestonespreventnormalexpansionofthenest．3ANET（1885）reported
anaerialqueennestof Vc和brvwithtwoenvelopesheets．Thesefactsshowthatthisspecieshastheability
tochangethetypeofenvelopeaccordingtoitsnestsiteenvironment，though Vc7Vbrvj7dvqfbsciatainJapan
alwa）．Sbuildsasphericalenvelopeeitherinasubterraneanorinanaerialnest．Themanycompleteenvelopes
inqueennestsisrnoreadvantageousasaprotectionfbrextrernetemperaturenuctuationsintemperatereg10nS．
Inthisrespectthernany−Sheetedenvelopeof Vsimit〟mq，aSWellasthatof挽甲uねandDolichove甲u毎
isconsideredprincipallyanadaptivefeatureagainstcoldsprlngWeather．Itservestotrapairinspacesto
greatlyminimizeheatlossduringthesolitaryperiodwhenonlythequeenandafewmaturelarvaeproduce
heat（IsHÅY19？3，G柑01977）．
Inthesetwospeciesanotherdramaticstrategyinnestinghabitsistheadoptionoftherelocationofnests．
Thequeensofbothspeciesbuildprimarilyinsmauspacesornarrowcavitieswherethemicroclimateismost
Suitablefbrtheearlystagesofdevelopment，aStheabilityofthecolonytoregulatetonesttemperatureis
SeVerelylimited．Whenthenestbecornestoolarge，thewholecoIonymovestoanew，mOreSpaCioussite，
becausethecolonyisthenabletoregulatethetemperaturewithinthenest．Thus，thishabitalsoappears
tobeanadaptationtocoldweatherbyproducingworkersasrapidlyaspossible．
Thermoregulationshouldbemuch easierformaturecoloniesthan fortheyoungeronesofthe＼・eSpine
WaSPSbecausethetemperatureinsidethenestisefftctivelycontrolled，aSthecombsaresurroundedbynumerous
Smauairpocketswiththeincreaseinnumbersofenve10PeSCOVerlngthecombs．Thisabilitytothermoregulate
enabledthevespmewaspslntemperatereg10nStOmakebetteruseofthe10ngerSeaSOnOffavourableweather
forthedevelopmentofthecolonyandthehigh，relativelyconstanttemperatureofmaturecoloniescanpiay
avitalpartinqueenproductioninthecoolautumnseason．ComparedtothevesplneWaSPS，pOlistinewasps，
Whichbuildanestwithoutanenvelopeamuchshorterlifecycleinwhichtorearthesexualsduringthehot
SummerSeaSOn（YosH∫KAW＾1962，WEST−EBERHARD1969，YAMÅNE1969，M＾TSUURA1980）becausethecoト
Oniesareunabletoinsulatethemselvesagainstunfavourableautumnweather．Thus，theproIongedlifecyele
OfthevesplneWaSPSintemperatezonesiscompensatedbynotonlyproducingmorequeensandmales，but
also building upmore populouscolonies which evoIvedefbnses against predators．
ⅠⅠⅠ．FEEI）ING HABITS
Thefbodofhornetsconsistsoftwowelldefinedclasses，1iquidsandsolids．Theformeriscarbohydratefbod
COnSumedmainlybyadulthorrletS，Whilethelatterisproteinfbodeatenentirelybylarvae．Thesesubstances
areobtainedbythehomets倉ommanydifferentsources，andtheyhaveevoIvedvariousstrategiesof蔦ndingthem．
Thischapterdescribesthe feedinghabits of掩岬aand some specificdifferencesin themodeoffood
40
Makoto MATSUUR＾
COllectionamongthem．
1．R）Od sources
l）Carbohydrates
Carbohydratesareobtainedftomtreesap，honeydewofaphidsandpsyllids，nOWerneCtarandothersources
aslistedinThble12．ThemostimportantcarbohydratesourceforalltheI包5PaSpeCiesisthetreesap
exudingfromlivingtrees，eSpeCially Quenusspp．（Fig．30）．Therelativefrequencyofvisitstotrees
Fig．30．Uninseminatedqueen（top），andworkers（Centre）ofTk5Pamandarinia，andqueensofI句PatrPPica（right
andbottom），Visitingtreesap．
sourcesbyvariousI句paspeciesisinthefb1lowingorder：mandarinia＞cnlbro＞analLs＞tTtPLCa＞simiHima・
ThetreeapertureexudingsaplSuSuallyoccupiedby Vmandarinia，Whichclaimtheterritorialrightby
chasingallthevisitorsofsameordifferentspeciesofhornets，beetles，butterflies，fliesandotherinsects
fromtheplacewherethesecretionisabundant・Bothintra−andinterspecifical1yadistinctlineardominance
orderisrecognizedamongthehornetsvisitingtreesapandtwoormorespeciescannotsettleatonetree
apertureatthesametimeasdiscussedlater・
FlowernectarisgatheredftomeightspeciesoffivefamiliesofnowerlngplantsukeCbmelliaspp・，C卸rutia
jqponica（Fig．31），MahoniajbrtuneLetc．，inwhichthenectariesarewe11exposedandtheflowershave
ratherabundantnectaries．Itseemslikelythat作中adonotrelyextensivelyonflowernectar，because
Comparativc bioIogyorlhe riヽ・eJapanescspeeiesorthcgenus tセspL7（H）7menOPtera，＼七spidae）
一泡blelZ．Carbohydratesourcesor物軍αSpeCies（1）
抱pβSpeCies＊＊
Species＊
SollrCeS
Sim Cra Ana MaIl ′Ro
Fam．Salicaceae
l．ぶ〝放gr〝ぐf胎伊ねMlq．
Fam．F喝aCeae
2．g〟g汀〟∫∫er柑′αTI王UN臥
3．9α川痛血mCARRAT11■
4．￠g／α〟甜THUN臥
5．朗蘭＝制御肋軌MÅKINO
¶ee sap
Fam．Ulmaceae
6．こ〟J〃〟∫卯rV的／∫αJACQ．
7．ぶe於ov〝ぷerJⅥ拍MAXINO
Fam．R11taCeae
8．Cか〟ぶ緑〃∫力如MÅRCOV．
Fam．Euphorbiaceae
9．肋／わJ〟∫ノ叩0〃た〟∫MuEL．ARO．
0
Fam．Berbcridaceae
lO，助力0〃ぬノb′J〟〃efFEDDE
0
Fam．Vitaceae
11．C町mJJ〝ノ叩0／lJ川G．ヽ（洒．
0
12．AJ，軍eわp∫ねわ佗痩7e血〃C打ね泡TRAN’rV・ 0
0
0
Fam．Theaceae
F10WerneCtar
13．C（”Jビ／仙ノ叩0′J／川L．
14．C∫〝甜明〟αTHUN臥
Fam．Urnbellirerae
15．掩錯加血椚棚卸網M血，
16．月〃gだ／加即妙cぬ（ねぎRANCH．
0
0
0
0
0
0
0
Fam．Oentianaceae
17．0だ〃′ね〃α∫亡α∂JVBuNG王；
く〉
Fam．Moraceae
l乱ダね録∫C即∫ピαL．
0
0
0
0
0
0
0
0
0
0
0
0
0
Fanl．AtlOna亡eae
19．月裏J〃f〃αか′Jo∂αDuAL
ぬm．Rosaceae
20．P川J川∫pe√∫血B人■＝引∵‖．
之1．㌣yr〟ぶ∫だrO擁αRE】ヰDER
0
Fam．Vitaceae
Riperruit
22．粕血油榔開L．
FaIll．Ebenaceae
23．か∼α勒ノ膵∫如たfTHUNB．
0
Fam．Solanaceae
24．みC叩e持′co〝e∫C〟／e〝J打招MILL．
Fam．Cucurbitaceae
25．C∼け〟／／〃∫わ〟J如月FoRSKA】‥
Fanl．BroIlleliaceae
26．月〃〃α∫ぐ0沼0∫以∫MERR．
0
41
42
Makoto MATSUURA
Table12．continued（2）
掩岬αSpeCies…
Sources
Species●
Sim Cra Ana Man l†0
Insecta
Fam，Coccidae
27．Cど′qp血ぎ／錯r〟如〃∫MASKELL
Fam．Psyllidae
Honeydew
28．顆〃α′0わぬeMIYATAKE
Fam．Aphididae
29．エβぐ如〟∫け叩ね〃胎GooT
30．乃ズ叩′e′αeかfごf血∫KIR餌LDY
O
3l．丁（扉血ⅣGooT
Mushroom
Oasteramyctes
Hymenogastrales
3Z．励み町′α血〃如0〃加IMAletKAWÅM．
Manred
33．Syru。jul。e
O
O
0
0
0
。
0
0
＊ScientincnamesofplantspeciesfromMAKINO’snewillustratednoraofJapanbyMÅXINO，HÅRÅandTsuY＾MA196i．
＋．Sim：l／：simil／i117a．Cra：lてcrabrq Ana：lてa170Iis．Man：レ：mondorInia．TTo：トて（rOPica．
F壬g．31．掩軍〟∫血fJ〃椚αWOrkertakingnect8r鉦omthe Fig・㍑・挽研削鞘彿ぬ血涙workerfbedi喝Onarig・
flow8rSOr Cり′用地ノ叩0〃加．
theyhaveabroadheadandshorttorlgueandprobablybecausetheamountofnectarco11ectedperflower
isso smallthatitdoesnot suffice for their nutritionalrequirements．
Thejuicesofwelトripened什uitsarealsoconsumedingreatquantitieswheneveravailable・Figsareespeciai−
1ypre飴rredbyworkersofmost掩軍αSpeCiesduri喝1atesummerandautumn（Fig・32）・Theattackusua11y
occurswhenthefruitsarerlpeOnthetreeandjustenou＄hfbrpicking．Theworkersofシ：mandaT．
frequentlychewandsuckthetissuesenmassandcauseseriousdamageandlosstothefruitgrower・W）rkers
of VlrppICaalsosuckthejuicesfromover・rlpe加itsofpeaches，pearSandper＄imrnonsattachedto
thetree．Incaseofgrapes，tOmatOeS，plneapPlesandwatermelons，Ihaveobservedsome掩軍aWOrkers
fbedingondroppedordiscardedonesonthegroundintheorchards・
Comparativebiolo＄yOfthefiveJapanesespeciesofthegenus物a（Hymenoptera，鳩spidae）
43
Fig・33・tセspa＼＼′Orkersftcdingonmushrooms：A．11L7／JaIi∫；B，Iくma17dariJJia（10p）and Lて（rOPica（boltOm）．
ThemushroomisanunusuallysourceorcarbohydrateぬrvesplneWaSpS．HoweveちWOrkersormost
Iセ軍αSpeCiesmakeitar喝ularpartoftheirdietinautumn（Fig．33）．Furthermore，malesoデH椚α〃−
dariniaappeartobemostattractedandtakeitastheirprlnCIPalfbodsourceafterleavingtheirnest．
1ha＼′ene、′erObser、′edthcm feedingonanyother foodsinthefield．
Honeydewproducedbyaphids，SCaleinsectsandjumplngPlantlice，WaSCOllectedfromleavesbyonly
Makoto MATSUURA
44
Vana］L5；Whileother挽研aSpeCiesscarcelyvisitedthesweetsecretion，thoughitiswell・knownthat抱卿壷
andDo（ic770、，eSPILlospeCiesfrequentlycollectilasarichsourceofcarbohydrate（DuNCAN1939，El）＼＼■＾RDS
l粥0）．
SyrupjulCeWaSCO11ectedwheneveravailablebymost掩軍aSPeCies，Whichhavebeenobservedlicking
itatapileofemptytinsand bottles．
2）Protein
Mostoftheproteinfbodco11ectedby掩spaspeciesconsistsofinsectsandspidersaslistedinrrhble13．
Judgingfrorntheserecords，thoughsti11insufficientinnumbersobserved，itseemsthatpreypreference
O‖頗卯di摘rscbaracteristicallywithspecies．
V：simiI（imotendstohuntawideran各eOfinsectsandspiders（Figs．34and35）．Morethan44species
Ofwingedinsectfrom8differentordersandseveralspeciesofspiderswereobservedasitsprey．Ofthese，
niesofthefollowingfamiiiesareparticularlypreferred：7abanidae，Syrphidae，Muscidae，Anthomyiidae
Fig．34．レi・5Pu∫imilIj111a、VOrkerhunlingalong−hornedgrasshopper・
潜血鞄
Fig．35．Preyloadsbrou如1tbacktothenestby挽野侶由血勒肌WOrkers一
Comparativebio王ogyofthefiveJapanesespeciesofthegenus ya5Pa（Hymenoptera，＼屯spidae）
Thblc13．Prcyo（拍spaspccics（l）
Prey species
Stageofprey＊
Frequencyofobser．
H∫∫J扉沼所々
Insecta
Orthoptera
Fam．Loeustidae
l．撤／叩￠d貪／〃〃∽沈αゐBoLIVAR
2．月細CJO椚0草加如（お〟BoLIVÅR
L．A
A
8
12
Fanl．’Ttttigoniidae
3．♪加〃gr叩Jどmノ抽αJαPoDA
L．A
9
4．月Sp．
L．A
8
Mantodea
托m．Mantidae
5‥撤憫確那感肌＝雨現隠侮STOLL
良（h．sp．
Odonata
Fam．Libellulidae
7．0′J如′r〟研けね〝g〟ぬ躇SELYS
8．旦y椚卵汀〟椚ノね留〟銅∫SELYS
9．Gn．5pP．
Hemiptera
Iもm−Cicadidae
l乱G殉紬聯血梱れ鴫叩ル甜摘MoTSCHLSRY
l王．タ由りや／紺相加g椚メ痴rfFABRICIUS
12．Aグe／刑〟〃β叩α′締柑WALXER
Lepidoptcra
Fanl．Noctuidae
13．Gn．spp．
Fam．Nymphalidae
14．掩〃e∬打f〃dねβHlミRBST
Coleoptera
Fam．Searabacidac
15，動画／／fαノ8pO〃血NEWMAM
Fam．Chrysomellidae
16．G11．SPP．
Fam．Attelabidae
17．肋co′力ね〟作〟血∫Ro王三LOFS
Hymenoptera
Fam．Vespidae
18．血J〃e／Jビ∫SPP．
A
1I
19．掩軍〟ねノ肋vfcep∫SトIITH
A
14
20．バクお〝7e／／娩用L．
A
＞100
2王．月．ぐgJ甘〃αFAIIRICIUS
A
24
Fam．Apidae
Diptera
Fam．Straliomyidae
22．タJecJ′c〟∫α〟′的rWALKER
ユ1．〟ビ〃JJe／血〃山川／J∫L．
45
Makoto MATSUURA
46，
′払ble13．contまnued（2）
Prey species
Stageorprey＊
Frequencyorobsen
Fam．Thbanidae
24．Gn．spp．
Fam．A5ilidae
25．C如e用d郎Sp．
27．C／？け∫0ねガ〟沼Spp．
A
28，gr加♂／由cピノでαJf∫FA】】RICIUS
29．エ〃′砂用p如加ん〃〟∫5p．
エリ
2
7
7
5
3nフ
3
00
00
■・▲
4
5
3
1 0ノ
■q
4 ■・1
5
射．Gn．spp．
A
Fam．Sarcophagidac
8
39．Cβ／／桓加）相加αCoquILLET
40．エ〝C∼／ぬぐ〃郎ぽrL．
A A
AA Å
3臥Gn，Spp．
Fam．Calliphoridae
A
37．劫／ね′e∫ねノゼ即ねFÅBRICIUS
A
36．Gn．spp．
Fam．Anthomyiidae
Å
35．肋∫ぐf〃〝Spp．
8
34．草y草加∼∫SPp．
Fam．MlユSCidae
A
33．勒蕗けqグ如加地〃拍deGEER
A
31．肋g〃即なぞ8乃β′αFABRICユUS
32．助椚er〟∫Spp．
A
30．grね撤ね〝ぴね′e〃仇rL．
2つJ81 7
A
Fam．Bombyllidac
26．Gn．spp．
Fanl．Syrphidac
42．β0だJ拍ぐ如r如ぽSp．
A
Fam．Scatophagidae
43．ぶぐα′力qp加rgぽ∫Je汀0用〟〟椚L．
Arachnida
Ararleae
44．GIl．Spp．
I‘：αJ柑JJ∫
Insecta
Orthoptera
Fa111．Locustidae
l．血相蛸朋明油日融如用oL王VER
Mantodea
馳m．Ma！ltidae
2．劫畑伽ぬ血相血埴肋STOLL
3−AⅣ〝削∽痛り知和ね加WÅSTWOOl）
Odonata
Fam，Libellulidae
4．紗叩舶W町旭押肌SEI−YS
25
ComparativebiologyofthefiveJapanesespeciesofthegentlS掩spa（Hymenoptera，Vespidae）
r払ble13．continued（3）
Prey species
StaさeOrprey＊
Frequencyorobsen
Hemiptera
ぬm．Cicadidae
5．A如∫研〟〃αqβα／娩用WÅLXER
Fam．Coreidac
6．尺jかOr拍∫ぐ／αVβ撤∫THUNBERG
7．Grl．5pp．
Lepidoptera
R旺m Sphingidae
乱軸力′〃ズ／fg〟∫けfBuTLER
Fam∴Noctuidae
9．Gn．spp．
Coleoptera
Fam．HarpaIidae
lO．C力／αe〃血∫Sp．
2
】1．Gn．spp．
Fanl．Scarabaeidae
i2．Gn．spp．
Fam．7tnebrionidae
13．タ／紺∼叩加地／椚〟∫〃JgrO伊α〃e〟∫MoTSCHOLSKY
3
18
Å
3
Hymenoptera
Iもm．Tiphiidae
14．7掬わぬspp．
Fam．Collelidae
15．Cβ地Je∫Spp．
Fam．Andrenidae
l軋。d細めⅥ別＝pp．
Fam．Halictidae
17．ガ〝′如〝∫SPp．
18．エ〝∫f￠gわ∬〝∽SPp．
Fam．Megac柚…dae
19．瀬窄αぐ力池謂〃J加）rか反YASUMÅTSU
et HIRASH】MA
20．A広路〟用ge〃∫ねCoc比良ELL
ユ1＝V．spp．
Ⅰもm．Åpidae
25．エα〝甘α椚〆ゆe／〃ぬSM汀H
26．ぶcg／わノ潤〃′J伽〟J乃SICXMANN
27．CerⅣrねspp，
Fam．Ⅵ≡Spidae
28．g〟椚ぞ〝ぞ∫SPp．
＝9．OIVIJCis（rOぐer．J∫dlt，ll・SeT］iS√＼USSURl三
30．ノわ／ノ∫け∫∫／ほ／／e〃／S∧しTSSしIRl三
つJ l▲
24．d暮Ce澗〃αFÅBRICIUS
Fam．Sphecidac
lノーJ つ▲
22．詑〃Ⅴね〃fα5p．
23．カタおJ〃g／／節用L−
47
48
Makoto MATSUURA
「mble13．continued（4）
Stagcorprey●
Frequenc〉rOrObscr．
A
3
Prey species
A
3
A
3l．RjqponiczISS入USSURLi
32．PI11a（1dari／川SSÅUSSURE
0
1
A
34．月cわ血g乃∫ねFÅBRJCIUS
﹁⊥
A
33．月ノαdwなαeDALLAToRRE
7
A
35．ノ和鱒pO少わぬJ乃dた〟SAUSSURE
A
つ︼
A
A
38．ダ′ec′′ぐまJ∫打〟r紳rWÅLKER
2
A
39．放〝〝eJぬ∫／／〟ぐe〃∫L．
1
A
FaIll．Asilidae
3 53エリ
37．物〟由ノ払vfαp∫SM汀H
Diptera
Fam，Stratiomyidae
長U
36．物α∫加′／／′椚βSMITH
1
1
40．C力￠g用d錯Sp．
41．Gm．sp．
‡もm．Syrphidae
1
2
5
1
A
A
43．旦y草力耶Spp．
3
1
A
4ユ．grJ∫拍／0／町血庸／JⅣ∫L．
45，肋∫ぐ〝SpP．
A
00
46．Grl．Spp．
A
2
47．Cα／／如舶用由泡Co（押ILLET
A
n7
44．On．spp．
Fam．Muscidae
48．エzJC〃ぬcαe∫〝rL．
A
Fam．Calliphoridae
A
50．βogJ′c毎rねぐ〝SP．
A
52．ぶぐ〝∼如炉如g（J∫Je汀0用r′〟沼L．
Araneae
l・：亡rロム′O
Insecta
Orthoptera
Fam．職t軸Oniidae
l．〃0／orノ】／0′〟SP．
Odonata
I屯m．Gomphidae
2．射通肋触触＝沼ね戚加SELYS
Hemiptera
‡もm．Ci（：adidae
3．CJγprOけ椚クα〝αノ叩0〃e那ねKATO
4．C／叩′0メガα／けぬ〃なrqル∫Cα拍MoTSCHLSKY
4 8 4
53．Gn．spp．
A
q′
Arachnida
A
0
51．GIl．Sp．
Fam．Scatophagidae
3
49．Gn．spp．
Fam．Sarcoph；唱idae
Comparativebio王ogyortheriveJapanesespeciesorthegenus物α（Hymenoptera，Ⅶspidae）
49
r払ble13．continued（5）
Pre）rSpeCies
Stageofprey．
Frequencyorobser．
5．♪／叫Ip／川和加ビ〃J尺／b′ノF八11RlぐrUS
6．〟ぞ血〟〃α叩α／推和WALKER
Hymenoptera
ml】1．Ⅵspidae
7．Po／J∫仔∫／，川／Jd（汀／／M∫SÅUSSURl三
Ⅰねm．Apidae
S‥力血／〃e〃枠用L．
托椚α〃（ねr加ぬ
Insecta
Mantodea
Qノ
‘V
3
L
L
4．物f〟0ガ〟／ノ描∫L．
2
l．撤佗′e〝Ode相打〟d的伽ST■OLL
2．創椚励血卯紬相加SERV】LE
3，On．spp．
Lepidoptera
Fam．papilionidae
uuA
I屯m．Mantidae
0
7
5
4
1
2
2
3
4
A
5
A
】0．JIJ〃Jほ血spp．
A
9．〟f′〃e／α埠）由〃de〃∫GYLLENIjAL
A
8．．4．spp．
A
7．月．〝撒q〆ね∫〝HopE
A
6．」／JO／J和仏ぐ岬r錯Hopl三
8
Fam．Sphir鳩idae
5．Gn．spp．
Coleoptera
鞄m．Scarabaeidae
3
】l．⊥〝ぐノ川0∫Je川〝SPP．
つ山
Hymenoptera
Fam．Apidae
0
5
L．P．A
0
L．P．A
13．＾．（1en711a F＾llR［ぐIUS
4
lユ‥■l両川血／昨′〝L．
17．柏叩α∫／J山〃血JαSト‖丁‖
L．P．A
】8．ト：c／Ⅶむ／■O L．
L．P．A
19．l：〟／J〟／止F∧bklぐ】US
L．P．A
20．rけq〆ぐ〝L．
L．P．A
ヱl，日刊川／〝ノTrハ・ナビ印）∫Sト‖T‖
L．P．A
鞄m．Argiopidae
22．勇将foクeα′乃Oe〃αAuDOUIN
1
Ararleae
1
Arachnida
3
L．P．A
．i ￠U
2 5￠U3 nフ
つム
1 7
L．P．Å
16．月r（）／／け呵l，JCÅMl三RON
7 2
L．P．A
15．月ノα〆けなαeDÅLLAToRRE
2 5
14．撤Jね招∫椚α乃d即∼〃打∫SAUSSURE
つム 4
Fam．Ⅵspidae
50
Makoto MATSUURA
Table13．continued（6）
Prey species
23．月．か〟e〃乃kカブfScopoLI
Stageorprey＋
Frequenc〉′Ofobscr．
L．A
25
lて什叩血
つム
3．月別♂〃d即f〃〟ざSAUSSURE
4
5
4つん2
4
P
＊L；はrva，P；p11pa，A；adult．
3
臥月sp．
P
7．撤叩卯かみねぎ〃蛮ぐⅣSAUSSURE
P
5．月代血澗〆CAMERON
6．月c力∫〃g乃∫ねFABR【CIUS
P
4．月ノαdwなαeDALLAToR鼠E
′0 1 25 8
3
0
2
P
l．伽／如g∫∫〃e〃e〃′SAUSSURE
2．書メ岬0〃fc〟∫SAUSSU鼠E
P
Insecta
Hymenoptera
Fam．Vespidae
3
P
P
L
L
L
L
L
L L L
Fig．36．挽甲βα〃α抽workerhuntingacicada，肋わ〃〟〃αq卯∼匝和．
andCalliphoridae．ThishornetisabまetocatchsuchactiveinsectsowlngtOitsveryswiftandagilemovements．
AfewlarvaeofanylnSeCthavebeenobservedtobecollectedbythisspecies，thoughmanysofトbodiedin−
SeCtS，Whichwereconvertedtomeatpellets，WereSOmetimescarriedintothenest．
Vana独alsotendstotakevariousinsects（Fig．36）．Morethan53speciesofinsectsfrom80rderswere
Observed，tOgetherwithsomeimmaturesofmantidsandsphinxmoths．Themostimportantpreywereadults
OfvariousspeciesofHymenopterasuchaspolistinewaspsandbees．Theworkersofthishornet，withits
agilenightactivity，huntparticularlyfbrsuchactiveinsectsvisitingflowers．
Compared to otherlセ5PaSpeCies，informationon pre），OfIてcIT7broare noIsurficient forconclusions．
Only8speciesofinsectsfrom40rderswereobservedandthedominantpreywereadultsofvariouscicadas．
AsalreadyreportedbyDAVIS（1925）thisspeciesisveryskillfulincapturingacicadaandtheworkersattack
ComparativebiologyorlheriveJapanesespeciesorthegenus物β（Hymenoptera，鴨5pidae）
Fig．37、Ibpqcrqbroworkerhun【ingacicada，Grap（OPSa／［riqnigrqf（J5Ca（a，
Fig．38．掩∫PaCrabroworkcrs handlingape11et oracicadabrough＝n the nest．
51
Makく）tO MATSUURÅ
52
Fig．39．柁岬a（rqPicoqueen attackinga Polistesjad両gqenesl．
Fig．40．Huntingor拉spalT7a11dqriniq：A，WOrkerattackingabeetleof′1no′〃E7IactLPrea；B，“，Orkcrcuuing
upthemantid．FlierodtL／opo（e／／昨m．
4speciesofcicadainandnearKibi．OneofthemostpreferredcicadasisCWtoLympanajbcialL；Which
isthelargestinbodysizeoftheJapanesecicadas（Figs．37and38）．Ifrequentlyobservedthatseveral
ofthecicadasweretakenoneafteranotherwhenveryabundantandwhentheircapturewaseasy．One
ObservationoncoIonyC7303withabout120workersshowedthat37loadsofonまythislargecicada，Which
WaSeStimatedtocorrespondtoatleast12individualsoftheadultcicadawhichhadbeenbroughtto
、thenestby5markedworkerswithinthreehour＄OnAugustlOth，1973．Inanothernest（C6802）withabout
240workerstheloadsofthecicadaoccupiedasmuchas71．6勒ofthe74loadsexaminedatthenest
entrance on August6th．1968．
VtrqpicadependsalmostexelusivelyonthebroodofPolistinaewaspsforitsproteinfood（Fig．39）．
王haveneverseenthishornetattackinganyotherspeciesofarthropod．A118speciesofpolistinaewasps
OCCurrlnginandnearKibiwereattackedbybothqueenandworkerhornetsofthisspecies．Adetailed
accountisgivenin the followingsection．
Vmandariniaisnotveryskillfu11atcapturlngagileinsects．Bothqueenandworkerhornetsofthis
specieshuntlargecaterpillarsandbeetles（Fig．40）aslistedinl抽Ie13．Butthemostremarkabletrait
Ofthishornetisthehuntingofothersocialwaspsandhoneybees，attaCkingandoftenexterminatingtheir
Comparativebio10gyOrtheriveJapane5eSpeCiesがthegenus掩箪α（Hymenoptera，Ⅵspidae）
53
COIonies・Thesehabitsarequitediffbrentfromthoseofother物aspeciesandthepeculiaritiesaredescribed
laterin thischapter．
21Fooddependenceo＝句卯（rOPicaon polistine＼＼′aSPS
掩spalrqpicadependsonvariouspolistinewaspsfbritsproteinfood・Thoughthehabitofattackingthe
lessersocialvespidsisreportedbymanyobservers（Rqpa［idia；RurrTER1916，Stenogaster；WILuAMS1919，
WARDl粥5，タロ／如e∫；So甑N1927，1951，SAKAOÅM！andFuKUSH王MÅ1957，撤／叩0か抽；VECHT1957，SEKⅢMA
etal・1981），thesepreviousdescriptionsaremostlyfragrnentaryorsuperficial，andthespecificこharac呼Of
thisspeciesisnotwelトknownindetaiI．Thissectiondealswiththehabit，Whichisquitedifferentfromthat
OrOthercongenericspecies．
1）Collection ofprey
When Vt7WECaattaCksanactivenestofFbHsteswaspsthemajorityoffb触esworkersescapetoplaces
nearthenestsuchaseavesofbuildingsortwigs，leavlngthequeenandafewworkersonthecombwith
noslgnOrreSistance・Thehornetpayslittleattentiontosomeworkersstillsittingonthecombandopens
firstacocoonwithitsmandiblespullingouttheprepupaoryoungwhitishpupa．Sheholdsontothe
COmbwiththetwohindpairsoflegs，keepingthepreyinthefbrelegsandmandibles．Then，Shebegins
tomalaxateitthorouめ1y，turnlngitcounteトC重ockwi5eWiththe如elegs．Sまmultaneously5hesucksrluid
fromthepreyandstoresitinhercrop．Afterthecontentsares＼＼・a1lo＼Ved，thevictimismadeintoasma11
ballcomposedofunconsumedremnantsandisthrowntothegroundundertbenest．Inthecaseoflarge
Po／istesspeciessuchasRjadwigaeandRrothnqyちaSlngleprepupaprovidesenoughfluidtobecarried
away，butinsmallerspeciessuchasRsne他niandRchinensねthecontentsoftwoorthreeindividuals
aresuccessivelyswallowed．Thcdurationspentinchewingwas130−250sec．（声ニ185．0，n＝72）．
Thepreyweretransportedasfluidin34Oobservationsexceptforonlytwoinstancesinwhichthehornet
transportedthefoodintothenestbetweenitsmandiblesasaslnglefreshballinasimilarmannerto
thatorother物αSpeCies．
ThePoILstesadultshavingescapedfromtheattackednestreturnafterthehornetfIiesaway，though
thenestoncefbundbythehornetiscontinouslyvisitedbytheindividuaIforseveraldays（Thble16and
Fig−42）・A丘eralltheprepupaearetaken，thehornetsshiftstothefullgrownlarvaeorolderpupaeand
finauytolaterstagepupaeandimmaturelarvae．However，fhaveneverobservedthehornetcatchany
Po／isIesadults．
Occasまonally，alienhornetsappearsnearthenestduringthechewing，buttheyarechasedawaybythe
“owners”withorwithoutcombat．lnallcasessofarobserved（＞400），jblistesnestswerealwaysattacked
andoccupiedbyasinglehornet，thoughSAXAGAMlandFuKUSl寸iMA（1957）statethatthehornetworker
Oftencomesbackaccompaniedwithoneortwonestmatesafterthe firstvisit．
2）Phenology ofvisitsto PoIis（eSCOlonies
OverwinteredqtleenSOf魚JisleswaspsbegintoestablishtheirnestsinearlyNrniddleAprilinandnear
Kibi．In掬ILstesjadwなaちthemostpopulousspeciesinthearea，thedaysofthefirstfbundationoftheir
nestwere：Apri110th（1967），15th（‘68），13th（‘69），15th（‘70），2nd（‘71），7th（‘73），13th（＜74）and13th（ワ5）．
Thereafter，developmentoftheRjadwなaecoIonyproceedsgenera11yinthefbllowlngSeaSOnalpattern
（Fig．41）：（1）Emergenceoffirstworkers−earlyJune，（2）Reproductionofmalesandnewqueens−mid
・JulytolateAugust，（3）Maximumdevelopmentofbroodpopulation−1ateJulytoearlyAugust，（4）Rapid
CeSSationofbroodrearing−lateAugust，and（5）Disintegration−mid−tOlateSeptember．Ontheother
hand，V．trqpjcaqueenscommencetobuildtheirnestsinlateMaytoearlyJune（Fig．2），abouttwo
monthslaterthanPoぬtesspecies，justwhenthefirstfewworkersofPolisteswaspsbegintoemerge
in thecolony．
Thefirstattacksby Vt／叩jcaqueensonPol加escoloniesareobservedfromearlytomidJune（徽ble
14）．Visitsbecomeconstant，therea鮎ちWiththecoIonydevelopmentthroughoutthesolitarynestingperiod．
AftertheemergenceofIて（ropicaworkersinmid−tOlateJuly（1もble3andFig．41），thequeenstillcon−
tinuesfbraglngaCtivitiesuntillateJulytomidrAugust，thenceasestoattackjblistescolonies．Onthe
Makoto MATSUURA
54
雄
0
O
㈱
藁葺麦細山 g 慧一︶ S J ﹂氾 U
■U
8
白山＞∝山S 斡 O
SトS山芝
O
蘭
山〇．〇芝
′0
0
8
。
4
J〇．〇之等山∑
O
4
Fig．4J．Scasonalchangesinthenumberoraclivenes【SOrFb／is［esjadttligoe（OpenareaS）andthoseorthenests
attackedby Vaspqtrppica（hatchedareas）intheⅥなkayamaFruitrrteeExperimenta！St．fieldatKibi
in197l．Solidcircles；themeannumberofcellsin撤Hstesjadwigaecolor虚s：Opencircles；themean
numberoflarvaeandpupaeinthecoloniesofRjadwigae．
lbble14．Recordsorthc first and finalattack byレさ5PaIrOPicaon polislinccolonies jn and near Kibi，1966−1975．
Year
’66
，67
，68
，69
り0
り1
り2
，73
’74
り5
Bylrて（rOPicq First attack Jl））l．10Jun．ユJun．】Jun．JJun．1JJun．3Jun．ニJun．5Jull．3Jull．6JunJ17
queens
Finalattack ワ Ju】．30 ？ Aug．2Jul．22 Aug．16 ？ Jul，18 Aug．5Jul．25Jul，30．1
By LJ：tropica First attack Jul．ニ4Jul．ユO Jul．コ5JuL15Julユ5JuL．：5Juu3Jul・ユ6Jul・：O Jul．22Jullユニ・5
WOrker5
Finalattack ？ Aug．24 Aug．2＄ A喝．25 ？ Sep．7 Sep．2 Au払26 Aug，28 Aug−25A喝．28．4
Otherband，WOrkersofVtrppicabegintovisitthenestsftomlateJulyandcontinuetoattackthemuntil
lateAugusttoearlySep【ember（Tbble14andFig．41）．In【てtropicacoloniesmalesandnewqueensare
rearedfromearlyJulyandemergeinlateAugusttomid−SeptemberandtherearlngOfthelarvaeordinari−
1yceasesinmidYtOlateAugust（Fig．77）inparallelwiththecompletionofbroodrearingin撤触escoト
Onies（Fig．41）．Atthetimeofemergenceofthesexualsfbragingacti＼′ityof、、′Orkerschangesfromprotein
foodtocarbohydratessuchastreesapandfruitjuices．TheworkersdiscontinuetherearlngOfthere−
mainlnglarvaebeforethedearthofproteinfbodsources．Sotheymakenomorevisitstothepolistine
nestsafterrnid−Septemberandstopfbedingthelarvaeinthenest．Thusitisclearthatthelifecycleof
Iく／ropicaiss）・nChronized≠・ith thatofpolistinewasps，theprincipalfood source．
3）RatiooffbIistesjadwigaecoloniesattacked by V（rqPica
rrable15showsthenumberofRjadw如enestsattackedby Vtrqpicq，andtheperiodinwhichtheywere
destroyedwithintheconfinesofthefarmsteadofWakayamaFruit7ieeExperimentalStationoverca．30
hectarearea，1969∼1975．Thenumberofnestsobservedduringthetimefromnest foundingto first
ernerBenCeOfworkerswas246，Ofwhichonlyonenestwasattackedduringtheseven−yearperiod．After
WOrkeremergencethenumberofnestsattackedincreasedmarkedly，thoughtheattackedratiowasrather
Variablefbryearandperiod．ItrangedfromlO．9甲￠in1970to93．2勒in1973duringtheperiodfrom
emergenceoffirstworkerstofirstemergenceofsexuals，and12．5巧もin1970to80．0％in1971duringthe
ComparativebioIogyofthefiveJapanesespeciesofthegenus掩spG（Hymenoptera，Vespidae）
55
7hble15・Numberandphaseof馳加esjddw如enestsattackedby抱patropicawithinthcconfinesoftheⅥなkayama
lTruit TreeExp．Sl．．1969・1975．
’鞄ar
【
lI
Tbtalno．
nests through
Periodsl−III
11l
1969
∵∵∴∴∵ユ∴
No．nests auacked No．ncsts attacked No．11CSIS auaCked
／observed（勒）／observed（勒）／ob5erVed（％）
0／25
（0）
1970
0／27
（0）
1971
0／88
（0）
0／18
1973
1／48
1974
0／22
（0）
王7／45
（37，8）
7／58
（lヱ．1）
20／121
（16．5）
25／47
（5ヨ．2）
45／58
（77．6）
（2．1）
16／39
（41．0）
（0）
1タフ5
0／18
14／30
（46．7）
（0）
Tbtal
or
トて什Op血
Observed（％）
■■
1972
attacked／
No．nests…
l／246
90パタ8
（45．5）
（0・4）
53／i13
144／398
（46．9）
21
（36．2）
＊Ⅰ：befbretheemergenceoffirstworkers，Ii：thereaftertilltheemergenceofsexuals，ilI：thereaftertilldisinte＄ration．
＊＊Nestsdiscoveredwithinabout500m汁omtheⅥねkayamaFruit甘eeExp．St‥
3
0
S之01ト＜＞∝山S寧○
0
2
ト聖U
■よ￥
1 2 3 4 5 6 7 8 9 川
DURATlON（DAYS〉
S之0︼ト哺＞∝山S蹟0 ﹂〇．〇こ
1 2 3 4 5 6 7 8 9 川】1 ほ
DURATIONS（DAYS）
Fig．42．Predationperiodof掩5Pa（rqpLcaononenestofPolis（esjadwなaebyaqueen（upper）andaworker
（lo＼＼・er），1969−1975．
56
Makoto MATSUU見入
rrhble16．Somerecords ofattack byqueensand、、′Orkers orlセspa（rOPicL70n Polis（eSjod両g（7enCStSin1973．
Nest codc
Or
No．or
H什叩J甜
Cells♯
No，■
rirslattaCk
No．o（
4
3
5
︻J
0
3
■j
つ︼
0
′0
5
l
3
つJ
つJ
−．1
￠U
l
9U
tl
つ一
−一
・1
っJ
．．ま
つ一
1
つJ
−1
3
7
25
Jul．之9
2
ハブ
5
4
29
Aug．Z
jO
Jul．27
27
Jun．26
3
封
Jし1Il．コ：
4
20
Ju†l．25
Juil．6
eaten／day
nノ
Jun．18
by HJ．＊…
つ︼
ほ
Jtln．i7
u。
3
Jun．13
（拍days）
13
封
PjユO19
Pj 3026
Pj 3027
Pj302＄
Pj 3035
Pj 3041
Pj 3047
Jun．9
貢naiatlaCk
JJu
謂
Pj 3019
Pj 3017
01 01 01 銅山揖 02 那 01
q q q q q q q q W W W
7
2
Pj301ユ
l）uraLion o［ lbtaI］10．0［
attack brood eaterl broo（】
月カか鴫朋
Pj 3007
DaLe Or
4
︵U
3
8
02
●the numberorce11sin F！jodll・igaenest on theday＼＼，henlhe rirslaltaCk byト：TrOPicE7、、・aS Observed．
頼q：queemhornet，W：WOrkerhor11et．
＊＝Thenumberexciuding痛oseoreggsandトⅠ王Iinstarはrvae．
03
periodafteremergenceofsexuals．JudgingfromThbles14，15andFig．41，tOgether、＼・ithotherobser＼′a−
tions，itseemslikelythatattackbefbreemergenceofPo飴tesworkersisoniyperfbrmed by VtrqpICa
queensandthoseafteremergenceofthesexualsaredonebytheworkerslargelyinlateJulytomid血gust
WhenthedevelopmentofbroodinthePoHstescoIoniesreachesapeak．
Ontheotherhand，thenumberof V（rOPicanestsdiscoveredwithinabout500mfromtheExperimen−
talStationvaried fromonein1970to fivein1971duringthetheseven−yearPeriod（7bble15）．1n1970
theattackedratiowasremarkablylowiJlthecomparisontothoseintheothersixyears，eitherbefbre
Orafteremergenceofsexualsinthe撤Iistescolonies．Thismaybeduetoalowdensityof V trqptca
around thesur＼・ey areain this year，
4）Predation period on a jb／Ls（e∫neSt
Oncea［／：（ropicafindaFb／is（eSneStShevisitsthesamenestdailyuntilallthebrooda、・ailableareex−
hausted．QueensofVtrqpicaconsecutivelyattackedanestofRjadwigaefbrl∼9dayswithanaverage
Of4．5days（S．d．土l．6，n＝101），andtheworkerstookl∼12dayswithanaverageof7．2days（S．d．土2，4，
n＝75）topreyuponacolony（Fig．42）．Thediffbrencebetweenpredationperiodsofqueensandofworkers
isprobablyduetothedifferenceincolonysizeofRjadw＊aeatthetimewhentheyareattackedby V
trqpica．Theattackby Vtropjcaqueenstakesplacemainlyontheco10nyattheperiodofslowincrease
＼Vithasmallnumberofbroodavailableforpre），（Fig．41），but＼＼′Orkerspreyedupon8．8−13．3irnmatures
Perdayjn】ateJulytoearlyAugust（7hble16），becausemorefoodisthenrequiredtornaintaintheircoト
OnyaSit becomeslarger．
5）FateofattackedcolonyofPo／is（eS
Astheattackofト：［rOPicaisrepea【edunti】alZtheimma【ureStageSareCOnSumed，thenestsusuallycou】d
notrecover．WhenaPoHstesnestisattackedbeforeemergenceofsexuals，the＆dultwa5pSfinallyabanr
dontheirorlglnalnest，and89．1甲bofnestsattackedattheperiodbetweenemergenceofworkerstothat
Ofsexualswerereconstructedbytheworkersneartheoriginalnest（′払ble17）．
Completeobservationsfromthe＄tarttOthefinishofarelocationarescarceandthefouowingdescrlp−
tionismadebycombiningfragmentaryrecords．Fromthestartofattacksby Vtrqpicqthequeenand
WOrkersofPoHstesstoptheirbuildingactivlty，thoughtheycontinuetostayonthecomb．After Vtrqptca
haveattackedthecolonytwoorthreetimes，SOmeWOrkersleavethenestandflyabouttosearchforanother
nestingsitenearby．WherlaSuitablesiteisfbundnotfarfromtheoriginalnestthescou亡WOrkerssettle
ComparativebiologyorthenveJapa】1eSeSPeCiesorthegenus陀堺〝（Hymenoptera，ⅥさSpidae）
57
Tablc171TIlerateOrPoli5［c5jad”Iigaencstsat【aekcdbylセ5PatrOPicL7．1964・1975．
No．ornests
attacked
4
192
75
Reconstructionaモ Åbandonmentor Aさ即e脚ion
another site
2（50．0％）
171（89．1鴨）
（i59）（82．8甲0）…
8（10．7甲b）
181
disappearance
aroundlhe nest
Site
2（50．0勒）
21（10．9叫〕）
1（1，3％）
24
66（8臥0勒）
66
＊ See’払ble15．
…Numberornests reattacked byレ：J和〆Cαa触rreconstruction．
Ontheplaceandspendmostoftheirtimerestingonitwithoccasionalreturnstotheor聯nalnest．With
thestartofsearchingfbranewnestsite，theworkersintheorlglnalneststopbuildingactivitiesandthe
qucenalsostopsoviposition．Then，theyleavetosearchfbrthenewnestsite，Whichisusuallysituated
inashelteredplaceorinabush．Aboutadayafterthearrivalofthequeentheworkersusuailystart
buildinganewnestandthequeenb喝instolayeggs．
Thedistancebetweenthetwonestswasmostlywithinl∼3m，butreached4rnatthemaximum．After
theyreconstructanewnest，mOStOftheworkersfrequentlyvisittheorlglnalnestforseveraldays，though
theyneverreusetheabandonednestevenif VlrppLCaStOpattaCkingit．However，1590utOf171nests
（93，0勒），WhichhadbeerlreCOnStruCtedduringearlyJunetomidJuly，WerereattaCkedby Vt7PPicabefbre
theyproducednewqueensandmales．Ontheotherhand，afteremergenceofsexuals，88．0％oftotalnests
attackedby V t7VPicadid notconstructanewnest，andalltheadultwaspsremainedonornearthe
emptynest untildisintegrationin September．
3．Grouppredationby掩軍amandaJlniaonnestsofhoneybeesandsociaiwasps
王tiswelトknownthat抱卵用冊漉ぬ痛飢加ksbeehivesinmassandcausesseriousdamagetohoneybee
COloniesiIIJapan（OK＾t）＾1967，M∧TSUUtuandS∧…G＾NI1973）．Theattackb）Tレ：mandariniaarenotlimited
tothehoneybeebutaremadealsoonnestsorotherlessersocialwasps．Thehabitsofthishornetarequite
different丘omthosebyother挽軍aSpeCies，aSthefbrmercausescatastrophicdamagetothevictim’scolony
Whiletheattackbycongenericspeciesisrestrictedtoindividualhunting，withonlyminorefftcts．Inthis
SeCtionsomeofthepeculiaritiesofattacksby V mandariniaaredescribed．
1）Visitsby物αSpeCiestotheapiaries
Variousspeciesof Va5PaPreyuPOnhoneybeesandlessersocialwaspsastheirproteinfood（Thble13）．
Prable18showsthefrequencyofvisitstoapiariesbyfiveI卸aspecies，eStimatedfromthenurnberof
SpeCimenscapturedinhornettrapsatanaplaryinKibiforatenpyearperiod．Amongthespeciesvisiting
Llleapiary，V川andal−（rliaandトニsimi／Iimawcrefoundtobeimportant，andtheformerspecieswasover−
Whelmlnglymoreabundantthanthelatterone．However，therewasastrongtendencyfortheratioof
Visitsby VsimilZimatotheaplarytOincreaseincompensationwhenthoseby V：mandar−iniadecreased
insomeyears．Vsi］nillimarareiyfrequentedtheaplarieshavingbeenvisitedby VmGndarinia．Evenwhen
Visiting，theywerechasedawayorattackedby Vmandariniqwhichattemptedtocatch Vsimil［bnqand
irsuccessrul，tOOkitbackto thenest．
TherelativeabundanceofthesetwospeciesmayvaryaccordingtoIocality．InsouthernJapanboth
t（mandarinjaandlくsimi！limaseemratherabundantandthenumberofthelatterspeciesvisitingaplaries
rarelyexceedsthatofthefbrmer，butthesituationisfrequentlyreversedinnorthernJapanwhere Vman−
dariniaisregardedasless commonthan Vsimil［imちaSrePOrtedMATSUURAandSAKAGAMl（1973）．
Threeothel■SPeCies，lくana／is，［／：cJVblVandl／：（rOPico，WereSCarCe：Ofthese，レ：crabroandレ：oTlaIis
actualiyhuntedhoneybeesattheaplary，buttheywereofminorimportanceaspests．InthecaseofVfrqpica
5＄
Makoto MÅTSUURA
T汰ble18．Numberof掩spqspecieseapturedinhornettrapsattachedtohivcsatanaplaryinKibifbralOぅ′ear
Period．
No．or掩岬αSpeCies
Year
JI柑JI（ね「／／J旧
∫ル〃／／／／川β
No．
恥
町o No．
l
l
l
7・−−3
l
5
l
3
l
l
5
l
4
王
1
1．〇
0
4
1
1．L
0
0
O
0
O
L
O
O
O
1▲
′n︶
O
3
O
0
O
へJ
つー
7
L
つJ
O
lg74
5
■−1
】969
l
′0
8
7
1
QU
∧U
0
−
l
197】
l
4
くJ
2
7
′
．1
．〇
8・0
︵U
︵yU
5
一l
−
S﹂く⊃n︻＞−凸Zl
l
1
1966
′−U・4 4 Q′．4 5 7 7 ￠リ 4
3 ′0 7 っん ′0 3 8 ′0 0 つ▲
点㌧J︷Jハ〃nけ点ドJ＼βハ〃J
‘U
00
．
2
0ハY
0
八
0X︶
5
1
爪
りり
品 ∬6
0
2
1
くJ
1
●l，
1
4
1
つ︼
−1
9
a
Ⅶ
1
3
4
1
只︶
凸7
2
n7
つー
q′
2
5
只︶
9
Qノ
n7
8
12
5
2
7
6
8
2
n7
′O
3
Q′
O・■1
q′
9
10
′0
■J・J−
ー1
5
7
■・1
l
0
1J
′0
11
16
％
甲o No．
5
4
4
・・−−
0
3
8
7
n7
8
4
Qノ
︵y
0ノ
′0
7
只︶．J
7
‘U
Qノ
1
7
7
4
7
00
8
4
？
q′
OO
7
‖
21
8
1
10
No．
0
4
2
▲フ
ハy
ヰ
′hU
つJ
6
7
00粥
粥9696削る580鋸那70
00
3
′0
0 2
4 43 00 34−▲
5 ヰ34■リハ
ノ‘
2
VU7 5
′0
ハブ
′0
′h︶
0
7
甲b No．
Tbtal
nO．
什叩血
er抽√0
ロ／Jロ他
隻︶
rJ
O
∝叫讐≒≡
5
■U
︹J
5
5
0
5
′＿】一〝li】用70195＿J肌． ′／−ハ、へ．一＿−ノ＼′＝叫h 」ULIAGsEP】dcl㌶○−∨J’uLムG】sEPlocTNOV
一
一一−−，＋一・−・−■一・−一・−−一−−
‡
t
＿ノ′′、、、′、へ
】 ‡
Fig．43，Phenologyo＝句卯沼馴血血ぬworkers（脚）andother掩叩〝SpeCies（…−−−）capturedinthetraps
attachedto20hivesatanaplaryinKibiforaten−yearPeriod．Thehornetswcrecotlntedevcryfivedays・
someworkersoccasionallyvisitedtheaplaryarldwerccapturedinhornettraps，butsofarnoattackby
thisspecieshasbeenconfirmedthoughthisspeciesisregardedasaharmfulhornettohoneybeesinIndia
（SuLll】IÅH and M入日＾t）い・＾N1957）．
ComparativebioIogyofthefiveJapanesespeciesofthegenu5掩堺u（Hymenoptera，Vtspidae）
59
2）Phenolog）．Ofattack to apiaries
Visitsby挽軍aSPeCiestoaplariesexceptfbr VmandarinidcontinuedfromMaytoNovember；OCCaSiona11y
rnoreabundantinmid・Septemberbutnotformingadefinitepeakasin VmanddriTlia（Fig．43）．Incon−
trasttoother侮paspecies，Visitsby Vmandariniawereseasona11ywelldelimited．Noqueensorworkers
Visitedaplariesuntilmidl如1guSt．Thevisitstoapiariesbecarneconstantandgraduailyincreasedthereafter，
reachedapeakinlateSeptembertomid−OctobeちWiththenumberofdailyvisitsbeing3O∼50individuals
peraplary，Whichwasestimatedfmmthesumofthenumbersofhornetsdailyki11edandtrapped．Numbers
Of VmandarinjdworkersdeclinedsharplyafterlateOctoberinmostyears，andnoworkersappeared
inlateNovemberthoughtheystillcontinuedtheirnestingactivityJherapiddecreaseofvisitstotheaplary
iscloselyrelatedtotheemergenceofsexualsintheirnestsafterlateOctober；becauseworkersspendlabour
muchmoreincoliectingliquidfbodfbrthernalesandnewqueensthaninhuntingforthelarvae．
3）Phenologyofattackonothersoeialvespidspecies
A11theattacksonothersocialvespidspeciesby VmandaTlniawereobservedfromlateAugusttoearly
November（Thble19），andthepeakofpredatoryaeitivityoccurredinlateSeptenlbertomid−October，Whieh
Thblc19．Thetime and nunlberofnestsofsocialwaspsattacked by L／？5PalT7q11darint（7．1965−1975．
Jul．
Aug．
E h■1 L E
Oc【．
Tbtalno．
of nests
No＼・．
M L E M L E M L E M L attacked
′0
0
0
︵U
8
O
0
O
7
O
︵U
O
4
4
0
O
●
1
7
O
7
︵U
7
史U
0
11
nU
0
1
5
3
0
10
215
1
O
ハブ
18
O
0
1
1
4
8
12
拍
16
0
1
5
′0
O
︵U
八U
∧U
O
O
0 0 0 0 7 16 17 32 36 43 30 19 15 0 0
0
0
2
2
9
8
13
日
〇
0
0
3
0
0
8
0
1
0
︵U
0
︵U
2
2
0
︵U
n︶
O
7
1
O
0
O
︵U
O
1
4
0
O
︵U
O
Tbtalno．0ぎ
nests attaく：ked
O
挽p〟ぬノ肋ソた申∫
O
‡儀卿＝細画加
O
噸α∫勧′JJf如〃
掩叩閻椚描地
物那Ⅷ融珊
O
加地J錯5pp．
Sep．
＊Nestkeptatthevepiary．E，MandLmeanearly，mid−andlateineachmonth，reSPeCtively．
COincideswiththatofhoneybeesasmentionedabove．Ontheotherhand，thenumbersofworkersand
COmbsinvesplneWaSpSreaChmaximainSeptembertoOctober，andthelowercombscontainlarvaeand
PuPaeOfsexuals，Whichofferalargequantityofproteinfoodfor V：lT70T7dorinia．Probablythisrnaypart−
1〉・eXplaintheconcentratedpredationby V：ma17dariniaonhoneybeesandothersocialvespinecolonies
in these months．
However，Why VmandariniadoesnotvisitaplariesorothervesplneneStSuntilmidAugustisstillun・
SOlved．Itmayhavesomepre鈷rredpreywhichdecreaseinlatesummer；OrSOmequantitativechangeof
proteinrequlrementmayappearinparallelⅥ′仙thestartofrearimgsexuals．Alternatively；theattackon
aplariesandvesplneneStSmaybestartedonlyafterasufficientnurnberofworkershaveemerged．At
presentthereislittleconcreteevidencetotesttheseassumptions．
4）Predatoryhabits
Theattackby V：mandariniaisdividedintothreesuccessivephases，hunting，Slaughterandoccupation
both on honeybees and othcr＼・■eSpinenests．
a）Huntingphase
Inthecaseofbeehivesthebehaviourof Vmandariniainthehuntingphase主sbasicallythesameasthat
Ofl／：simiI／ilT7aandotherspecies，Whichattackthepreyonlybyindividualhunting，各ivingnoacuteand
CataStrOphicresuit．Maindifferencesare：
（1）Vmandar・ir2idworkersarelessskillftllincatch呈ngbeesinnight．Theyawaittheirchance，Sitting
nearhiveentrancesoronhivesmorepersistentiythanotherspecies．
60
Makoto MATSUU】iA
（2）Theyaremoreapttoseizebeeswiththemandibles．OncecounteトattaCkedandgnawedbyabee，
thehometfiiesaway鉦omthehive，remOVeSthebeewithitslegsintheairandimmediatelybitesittodeath・
In the hunting phase，therefbre，eaCh worker of V ma77dariniarepeats a chain perfbrmance of
visiting−Waiting−CatChingonebeeandpreparingameatba11ftomtheprey，smesosoma−Carrymgthe
ballbacktothenest．Atahivewbichcounter−attaCkweakly，2∼5hornetsmayvisitsynchromouslyor
alternately．Theyinvariablyrepeatthesameperformance，neVer＞leavlngthecorpsesofthebeesinthe
aplaryaSinthenextphase．Solongastheattackremainsinthehuntingphase，thedamagetohivesdoes
notexceedthatby Vsimil／imaverymuch，Orisoftenlessbecauseofthelowerfrequencyofvisitsreflec−
tingitssmallernestpopulation，andlessagilebehaviourthan H∫∼椚∼J伽払
Attacksonnestsofotherhornetspeciesalsostartwiththehuntingphase．TheworkerofVmandarinia
hoversaroundtheothervespmenestforquitealongtimeandshesuddenlypouncesuponanindividual
victimatornearthenestentrance．Afterfallingdownonthegroundincombat，thevictimisusually
ki11edwiththemandiblesandthemesosomaistransfbrmedtoameatballandcarriedbacktothenest，
aswithahoneybee．However，theattackedspecies，eSpeCially Vanalisand VcTVbrqoccasionallykills
the V：mandariniabycounteトattaCkingwithitssting．Inlargenestsof Vsimi［limaandIくcTyb7Vmany
workersusuauytaketotheairatthearrivalofシ：mandariniaandchaseawaytheinvadersothateven
thehunti喝phaseisonlyintermittentandseldomshiftstotheslaughterphase・Innestsofthesespecies
thataresmallduetodelayeddevelopmentororphanage，andnestsofVanalLfWhichneverreachalarge
size，thehuntingphasecan shirt toslaughter．
Thehuntingphasecancontinuefbranindefinitetimewithorwithoutshiftingtothenextphase・The
shifttotheslaughterphasepartlydependsonthedistancebetweentheaplaryOrthevesplneneStSand
the Vmandarir！ianest．Whenthevictim，snestislocatednearthe Vmandarinianestandreceivescons−
tantvisitsbyworkersofthesamenest，theshiftisnearlycertain，WhileatanaplaryOrneStremOtefrom
theoutskirtsofmountainsandhills，Wheremost Vmandarinianestsareconcentratedthefrequencyof
visitsislowandtheshifttothenextphaseisratherrare．
b）Slaughter phase
Oncetheattacksb的stothesecondphase，thebehaviourofレ∴椚甘m免佑納克workerschangesdrastica11y・
Inthecaseofbeehives，theydonotvisitdifftrenthivesalternatelyasinthehuntingphase，theyconcen−
trateattacksonaparticularhive（Fig．44），ignoringothersevenifplacedsidebyside・Ifonecaptures
Fig，44．Attacksby te5PamandartniaonahiveofApjsmel［昨7Yl：A，Slaughterphase；B，OCCupationphase・
ahornetatthisphaseandIiberatesitneartheapiary，Sheimmediatelyreturnestothevictimhivetorecom−
mencetheattack．Onethevictimhiveischosen，anOtherhiveisneversubstitutedthroughoutthisand
thenextphase．
Theattackinthisphaseisasimplerepetitionofslaughter・Usuailystaylngnearthehiveentrance，eaCh
hornetbitestodeatheachbeeapproachingherforcourlteトattaCk（Fig．45）．Thecorpsesareleftonthe
groundnearbywithoutbeingcarriedbacktothenest・Onceabeebitesherbody，thehornetoftensoars
up，remOVeSthebeeintheairwithitslegs，biteshertodeath，throwsdownhercorpseandreturnsto
ComparativcbioIogyofthefiveJapanesespeciesofthegenus掩SPa（Hymenoptera，Ⅶspidae）
61
Fig．45．［／bspE7mandcrinia、＼・Orkerbitingahone〉・bee．
thehiveentrancetorecommencetheslaughter．Theuseorthestingtokillbeeshassoねrneverbeen
Observed・Thenumberofhornetsattackingatthisphaseisalwaysmorethantwo，andupto50，although
theydonotworkcooperatively．TheslaughtercanbestartedfromearlymornlngtOeVenir唱，butusually
StartSintheforenoon．Onceslaughterdevelops，participatinghornetsdonotreturntothenestuntilthe
endofthephase．Or，Whenthephasecontinuestothenext，hornetsreturntotheirnestatsunsetbut
reattackthevictimhiveonthenextdayintheearlymornユng．
IndividualrnarkingshowsthathornetsparticipatinglnSlaughteratahivealwaysbelongtothesame
nest・Occasiona11yalienhornetsappearattheapiaryinthemidstofslau靡Iter，buttheyarechasedaway
by’legitimate’ownerswithorwithoutcombat．Ontheotherhand，neWlyarrivedhornetsofthesame
nestparticipatedirectlyinslaughter，Ornittingthehuntingphase．
Thedurationofslaughterisvariableaccordingtothenurnberofhornetparticipantsandtheintensity
Ofthedefense．Thedefensealwaysappearsduringtheinitialpartofthephaseandismostlylnitiated
bybeesofguardorfbragingages、、・hjleforagersreturni）1gtOthehi＼′eOftendonotparticipateinitsdefense
butleavethehivc．Whenmorethantenhornetsmaketheslaughter；COunteトattaCkssuddenlyceaseone
tosixhoursafterthestartofthephasewhenthepopulationofthevictimhivehasbeendecreasedcon−
Siderably．Manybeekeepershavebitterexperiencesofloslngtheirhivesduringtheirabsencefromaplaries
Ofonlyonehouroreven30minutes．Manysurvivlngbeesleavethehivebutsomeremainwithoutresistance．
Becausehornetsintheslaughterphasedonotreturntothenestnorchewthekilledbees，fboddeficiency
OCCurSaSSlaughtercontinues，reSultingintheappearenceoffrequentandpersistentfbodexchangeat
theaplary．Inextremecases，thehornetsstarvetodeathduringaproIongedslaughter．
Thenumberofbeeskilledbyhornetsvariesaccordingtothesizeofvictimcolonies，numberofhornets，
andintensityofcounter−a＝acks．1nsouthernJapan，thehi＼′eSmanagedbythenon−migratoryapiculture
SySteminvoIves15，000∼30，OOObeesinAugusttoOctober．Visitsby20∼30hornetsusuallyresultindeaths
Of5，000to25，000beesduringonetosixhours，unlesstheattackisartificiallyinterrupted．Hornetdeaths
duringtheslaughterphaseareusuallyonlyoneortwo．Inoneinstance，about25，OOOoutof30，OOO
62
MakotoMÅTSUURA
Fig・46・Attacksby挽御用矧感飾如“m植野肌面相肋は既旺Aパパ閥憫血幼 workers】）t曲1gOtllapupa
i）lLheocc叩ied ncst ol、l■：∫iJ11iIIim（I：B．shamblcsallLe）la L）nLl］e．
ComparativebiologyofthefiveJapanesespeciesofthegenus物a（Hymenoptera，Vespidae）
63
inhabitantswerekilledduringthreehoursby30horrletSWhileonlytwohornetswerekilledbybees．This
meansthateachhornetkiliedapproximatelyonebeeevery14secon血
Inthecaseofother掩5PaneStCOmbatinthisphasewasnaturallymuchrnoreseverethaninattacks
On4扉＝鵜朝沖澗（Fig・46，B）・Severalworkersofthevictimspecies（OrSOmetimesonlyonein巨α乃α胎）
COnn・OnteaCh Vmandariniainnightoronthenestentrance・Thetwoopponentsfhlltothegroundand
ri由tviolentlywitheachotbeちuSingmandible5，iegsandstings，Theslaughterphaselastsseveralhours
OrCOntinuestothenextday・Ina11sofarobservedcases，theslaughterphasealwaysproceedstotheoc−
Cupationphase．
InIセspu砲abouttenhornetscancompletelyconquerthevictまmnestduringonehourandoccupy
itwiththelossofonlyonetotwoindividuals・Adultvictimsaremostlykilledwiththemandibles，nOt
bystinging・Thecounter−attaCkby㍑j7avic叩SeemSnOtSOSeVereaSthatrnadebyApismeHifb7tt．Usually
fewdeadofbothpredatorandpreyarefoundatthenestentrance・Inanestoccupiedbyabout20V
mandariniaworker，SeVeraltensof VLj7dvjc甲SWereStillaliveinthenest，anddeparturesandreturns
Ofafbw㍑J侮Ⅴ∫c甲ざWOrkerswereobserved．Itisanopenquestionwbethertheslau紳terphaseishere
SOdistinct asinthecaseofthehoneybee．
Tbeeausalmechanismunderlyingtheshi魚抒omhuntingtoslaughterpha5eisasy飢unknown．As
theshiftusua11yappearsafterhornetshavereceivedintensecounter−attaCks，thelattermayreleasethe
Changeoftheirbehaviour．However，theshiftdevelopsonlywhenmorethanonehornetofthesamenest
particlpateSinhunting・Therefbre，itisplausiblethatsomeinterindividualrelationstimulatestheslaughter．
Moreover，thenumberofhornetsoftenincreasesinthecourseofslaughter．Somebeekeepersassumethat
theinfbrmationbroughtbyhominghornetsduringthehuntingphasefavoursthearrivalofothernest
mates・Howeveらthereisstilinopositiveevidencetocon蔦rmsuchanassumption．Astherecognition
Ofnestmatesexistsin Vmandarjniq，itisconceivablethatsomestimulantisemittedbytheindividual
Shiftingtoslaughterphase，Whichevokesaconcentrationoffurtherindividualstotheplace．Thefhct
thathornetsareattractedtotheindividualskilledandle魚atparticularpointsseemstofhvourthisassump−
tion，but theproblemisstillopen to criticalstudies．
C）Occupationphase
Afterthecessationofdefbnsebytheoccupantsandescapeofsurvivors，hornetssuccessivelyenterthe
Victimhiveornestandoccupyit．Theypaylittleattentiontosornebeessti11sumvlnginthecolony，Once
theslaughterphaseisattained，itinvariablyproceedstotheoccupationphaseunlessartificiallyinhibited．
Withinthehiveornesteachhornetwalksoverthecombs，andopensacocoonandpullsoutthepupa
（Fig．46，A）．Keepingitbetweenmandibles，ShefliesawaytoabranchnBartheapiaryornest，remOVeS
unnecessarybodypartsandcarriesbackthemesosomaofthepupatoherownnest．Atfirsttheypreftr
laterstagepupae，thengradua11yshifttoyoungeronesandfinallytolarvae，Whichareusuallydirectly
Carriedbackwithoutchewing．Soorlaftertheslaugherephase，thecorpsesofvictimsandtheirownspecies
arealsopreparedasmeatballsandtransportedtothenest．Butitceasesonthenextday，PrObablybecause
OrPutreねction．
エnthisway，theoccupiedhiveornestiscontinuouslyvisitedbytentoseveraltensofhornetsinthe
daytimeforseveraldaysuptotwoweeks．Oftensomeindividualspassthenightinthevietim，snest，nOt
returnlngtOtheirownne軋
Ilmandari11iaworkersintheoceupationphaseehangetheirdisposition．Theoccupiedhiveornest
isguardedbysomeindividualssittingattheentrance（Fig．44，B）．Theapproachofmen，dogs，Otherhornets
Ofdifferentspecies，andofthesamespeciesbelongingtoaliennestsoranyotherlargeanimalsreleases
attacksbytheseguards．Asinmostsocialinsectsawayfromnests，Vmanddri7midonotmakesuchaト
tacksonmenduringthehuntingandslaughterphases，SOthattheycanbebeatentodeathwithwooden
Sticks，etC．ataPはries．Intheoccupationphase，however，thisoperationbecornesdifficultbecauseofat−
tacksbyguards．Beforeattacking，eaChhornetoftenfliesaroundthemenoranyanimalsapproaching
theoccupiedhiveornestandthreatensthembyapeculiarloudclickingproducedbyrapidopenlngand
CIoslngOfthemandibles．ThisclickinglSprOducedinallJapanese物aspeciesatorneartheirnests，
Makoto MATSUURA
64
Fig．47．Distributionofvespinenestsandhoneybeehives（息）attackedby f＊amaT7darinia（◎）nearandat
theVhkayamaFruit取eeExperimentalSt．（WFTES）orchardandfbrestsin1972withthedateofthe
discoveryofmassattack．
冥加：物α∫f沼f沼椚び；Hα〝α：rα〃α胎；沼ノ払：掩甲捏由ノ加摘要方；月別：dp缶用e㈱m；血：d．α柑〃α
Thb】e20．Numbcr or＼・CSPincncsts attackcd bylセ5PuJ瑚11dal−i，Iiq，1965−1975．
No．11eStS
Observed
甲b of
1bar
（A）
Species
No．nests
attacked
nests
atta（：ked
つJ 7
つん 7
7
0 7
（y ′−U
8 ′n）
7 ′n−
′n▼ ′h−
）74 ’75 7btal
くー ′hY
（B）＊
5
4
1
0 2 4
n八︶
4
．〇
5
‘V
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〇
5 2
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7 0
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5 ∩︶
7
3
ユ
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0〇一lt−
3
￠U 5
0〇一▲U
1
4 0
1
7
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4
′0 3
3 つん
8
3 0
5 2
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4 0
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3 ′n﹀ 2
2
9
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3 5
3
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0 0
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4 0
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ト′ノ．．〃〝lイぐ叩∫
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′hU O
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7 5
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l∴血両／／J〃川
＊NumberofnestsobservedonlyduringtheattakingperiodfrommidAugusttoearlyNovember．
Comparativebio10gyOfthefiveJapanesespeciesofthegenus挽甲a（Hymenoptera，Vbspidae）
65
andisregardedasahabitcharacteristicoforprevalentinthegenus，‡n Vmandariniqitisproducednot
Onlyatnestsbutalsoatthehi、′eSOrne5tSOCCupiedandfrequently＼′isitedtreesapsources．Thisindicates
itswelldevelopedterritorialdefbnse．Therelativeaggressivenessofvariousspeciestomenatorneartheir
OWnneStSisordered：∽α〃（ね〟乃由＞頭如肋削ウ〝如何＞α習α飴＞打q練α．
AlthoughoccasionalentriesinlOhivesarerecordedfbrsomeotherhornetspeCies（lて卿nLsinFormosa
bySoN∧N1927；Vsp．inAfghanistanbyScHNEll）ERandDJ＾l“札1970）、alastingoccupationofvictimhives
ispreviouslyrecordedonlyin Vorienta駄FromthedescriptionbyIsHÅYetal．（1967），thepresenceofa
distinctslaughterphaseinthisspeciesisnotclearlItwouldbeinterestingtoobservehowVoTie細Zisdevelops
itsoccupationphase．Somebeekeepersassumethattheaimofhiveoccupationby Vman（ね血idistopillage
thestoredhone〉′lButasfarasmyobservationshavedetermined．hornetsdestroyhoneycombsbutrarely
takehoneytocarrybacktothenest．Asseen鉦omtheobservationsonvespinenests，thernainaimofoc−
Cupationmu5tbepredationonpupaeandlarvae．MostobservationscitedinMATSUURAandSÅXÅ0ÅMl（19T3）
OnOther rt5PL7SpeCiesconfirmthepreftrenceforthemuscularmesosomaoftheprey．Thisisinmarked
COntraSttOVisitsby掩軍uhlj7dvicqostoapiaries；thisspeciesseeksmainlythestoredhoney．F鼠EE（1970）
Showedthatmeso−andmetasomaofhoneybeesareequa11yprefbrredbytheEuropeanye11owjackets，物uhl
germanicaand VL v！J如血．ThismightindicatethatI々5PZLIaislesspredaciousthanレ旨5Pa．Asshown
in取ble12，H椚α〃（ねr飯ねworkersareespeciaIlyぬndortreesapbutdonot触quent恥wersexcept
forsomerarevisitsto CissusJqPOnicGandA〝軍eわクSisheterqpjo，Ha．Whendilutedhoneyandtreesap
aresimultaneouslyoffbred，Vmandarir7jainvariablyprefersthelatter．Thesefactssuggestthatthestored
honeydoesnotfbrmaprincipalobjectiveofthehiveoccupationinthecaseofattackonApLsme〟紳TV．
5）Fbraging distance
Theflightradiusoflて171a（1dariniajsonthea＼・eragel−2klll，reaChingamaxinlumOf8km．Thedistance
betweenanestof Vmandariniaandapiariesorvictim’snestsismostlyl−2km（max．2．5km）andserious
damageappearsmainlyatapiarieslocatedlessthanlkm鉦omhornetnests（KoJIMA1950，MATSUURAand
SAK＾GAMI1973）．Othervespinenestsattackedareaisomain1ylocatedlessthanlkm抒ornanestofVman−
da／’i11ia．Fig．47sho、、・SthedistributionofvespidnestsdestroyednearandaLtheⅥねkayamaFrui（Ti▲eeEx−
perimentalStation’sorchardsandfbrestsatKibiin1972．Of8nestsofvarious俺軍aand掩spuk7SpeCies
withinlkmftomanestof Vma77darin毎atotaiof5nest5WereattaCkedanddestroyedbyworkersof
thesamenestof Vmanda7血ia．AsfarasIknev％theworkersofthisnestexterminatedtwonestsof V
Simi〟imqtwoofVanalisandoneof挽甲uね」触vicqosduringSeptemberandOctober．Furthermore，they
attackedtwohivesofApねme〟推TaandonehiveofA．cerunawhichwerelocatedabout100maway鉦om
theirnest．Thesedatashowthat Vmandariniaplaysanimportantroleihcontrollingothervespidnests
in theneighbourhood ofトてmandarinianests．
6）Ratioofvespinenestsattackedanddestroyedby Vmanda7幼ia
Attackson掩叩LJ／aandIセ3PaneStShavesofhrbeenobservedonlyb〉・Vmaltdariniq．Thble20sho＼＼一Sthe
numberofnestswhichwereattackedanddestroyedbythehornetduringanlトyearperiodfrom1965to
】975．Theratioofnestsdestroyedby【てmL7ndal’i11iareached50．0甲0（34／68）ortotalnestsof Va5PZLkTJ7at・icqDS
Observedduringtheperiod打ommidAugusttoearlyNovemberinandnearKibi．Ontheotherhand，in
CaSeSOf挽甲E7SPeCies，thenestsdestroyedb〉′IJ：171a／JdarilZiawereonly4．帥0（2／40）inlイcrubrtl13．6甲0（16／l18）
in Vs加iHimaand15．4％（12／78）in Vana梅Ofthetotalnescsobservedduringtheattackingperiodof
7（mandarjnia．Largenestsofl物acanresisttheattackandinhibititattheincipientstageofthehunting
phaseirarelytheslaughterphase．Judging鉦omthedatashownin7hbles19and20，tOgetherwithother
Ot〉SCrVationsinthe員eld，therelativeresistiveabilityofvespidwaspsagalnSt V manddr飯jaisordered：
C／血ro＞SimiI／ima＞L71ta／Ls＞什jhl・icqoslthoughitmayvaryaccordingtosizesof＼′ictimcolonies，number
O＝加喝1nghornetsandintensltyOfcounter−attaCks．
AttacksorineStSOf Vtf叩icahaveso fhrnotbeencon蔦rmedundernaturalcondition．Thismaybe
duetotherelativerarityandshortannualcycleofthisspecies，Whichreachesthedissolutionphasebefore
theattacks by V marldari11iL7intensify．OnceilS neStSarediscovered and attacked，itis certain that this
mildesthornetmakingthesma11estnestswouldbeeasilyconqueredby Vmanda7勃ia．7h7OSmallnestsreared
Makoto MÅTSUURÅ
66
inmyvesplaryWereattaCkedandoccupiedeachbyonlythreeworkersofVmandariTtiaon19thSeptembeち
1973．Nointraspecificattackof Vmandar飯iGhassofhrbeenconfirmedeitherdirectlyorindirectlyby
thesuddenappearanceoデnumerousdeadnearthenests・
7）AttackonhivesoftheJapanesehoneybee，ApLsceru17a
Thefbregoingdescriptionshowsthatnestsofothersocialwaspsarevictimizedby VmanddT血jd・Only
largenestsof VsimiHimaand VctvbTVCanreSisttheattackandinhibititattheincipientstageofthe
huntingphase．Meanwhile，thereisanothersocialinsect，thebeeApLsceTUnaJqPO77icaFabricius，theeastern−
mostandlargestsub叩eleSOftheAsiatichoneybeewhichmaybeat払Cked・
SinceToKUDÅ（1924）itisweuknownthatthisspeciesavoidsthedamagebyretreatingwithinitsnestsat
thearrivalof Vmanddrinia．Closerobservationscon丘rmedthatA．certmadefヒndeditselfwe11againstthe
gianthornetnotonlybyapassiveretreatbutalsobyaparticularmethod・ThefollowlngObservationswere
madewithcolonieskeptinahiveadoptedinandnearKibi，aWOOdenboxwithseveralsrnauentranceholes
belo％Whichw鮎designedbyMitsuichiS姐A，the餌herofJapan彷eApic油u喝Who〕vedinAddanearKibi・
Atthearrivalorahornet，departuresたomthebiveabruptlycease．Theshimmerl喝，aWarnまngSO11nd
characteristicofthespecies（SAKAGAMI1960），isrepeatedlyemitted魚・Omthehive．Homingfbragerskeep
adistancefromthehornetandquicklyenterintothehivethroughholesremote鉦omtheenemy（Fig・48）・
Bythisrapidretreatwithnosoutarycounter・attaCk，thehornetusuauyrelinquishesherattemptandleaves・
Fig．48．BeginnlngOデthefbrmationoracircleor．4pおぐeJW〝〝amunda物α椚α乃ぬ血ねworkerattempti喝
toinvade the hive．
Irthehornetpersists，howeveちnume和uSbeesappear鉦omtheentrances托mOte打omit・Nevernyingnor
dartingsohtarilytothehornetasA．meH搾7Vdoes，SOmebeesapproachthehornet，directingtheirheads
totheenerny，ralSlngtheirmetasomaltipsandvibratingthewings・Thus，aCircleofbeesisformedaround
thehornet．Theattempttoseizebeesbythehornetseldomsucceedsfbrthemovementsofbeesareincorn−
parablymoreagilethantheirEuropeancousins・Meanwhile｝Onebeepouncesrapidlyupontheheadof
thehornet．Nearlysynchronouslyauothersinthecirclerushupontheenemyandotherbeesneartheen−
tranceconcentrateontheplace．AboutlO∼15secondsa鮎rtheattackbythefirstbee＞thehornetiscovered
thicklybyseveraltenstoseveralhundredsofbees・Shimmerlnginthehive，rePeatedsincethearrivalof
thehornet，CeaSeSanddepartureoffbragersrecommences・Thebeescoverlngthehornetkeeptheirpositions
Comparalivebiolog），Ofthe nYCJapanesespceiesorthcgenu5拉spa（H〉・rnenOPtera，Ⅵspidae）
67
fornlOrethan20min．wilhlittlemotion．Thesamemassdefenseisalsoapplied【Otheoccasionalhornet
thatentersahive．Thecorpseofthehornetshowsnoexternalinjury；Sheisseeminglykilledbystings．
Thus，theattackby Vmandarinぬisinterruptedattheinitialpartofthehuntingphase．Thehorneteither
iski11edorabandonstheattackwhichdoesnotdevelopintoitssubsequentphases．rhavehadtheoppoト
tunitytoobserveonlya食winstancesofthemassdefbnsedescribedabove．Butthelackofsufficientobser−
Vationsissimplyduetotherarityofcatastrophicdamageby Vmandark2血Accordingtotheexperience
Ofmeandseveralbeekeepers，lイma，ldarin（ane、′erVisitsthehivesofA，Cem［laPlacednearthoseofA．
mell締m．Amongabout300hivesandmorethantennaturalnestsofA．ce柑naObservedbymeintheareas
WhereA．me〝脾7Visalsokept，nOinstanceofslaughterorexterminationby Vmanddrinidhasbeenrecord−
ed．SeveralbeekeeperspossessingA．cemnahi＼′eS，includingone＼＼′hohaskeptseveraltensofcoloniesat
thesametime，invariablyrecordedtheabsenceofcatastrophicdarnageby‡てmandarinia．
Theefficiencyofbothattackanddefヒnsebetweenpredatorandpreymustbeenvisagedinitsequiiibrium，
Changingbysituations．IfacoIonyofA．ceTUnaiskeptinahivewithawideentrancespaceasusedfbr
A．me／I昨m，＼＼■hichmakesinvasionofseveralhornetspossible．theeffectivedefensedescribedabovemay
bedifficult．Oneinstanceofattackby Vmandd71niaonacoIonyofA．ceFmareSultinginmi1ddamage
WaSnOtedon22ndOctober1972．ThreeworkersofVmandariniawerenyingaroundin鉦ontofaA．cefYma
COlony，Which＼VaS keptinahi＼′e＼＼′ithanampleentrancespace．AFter ki11ingthesehornets，【hehive＼VaS
inspectedandthefb1lowingdeadwerediscovered：4hometsin鉦ontofhive，4withinthehiveandabout
80beesinfrontofthehive．Thehornets’nestwaslocatedaboutlOOmaway．DuringSeptemberandOc−
tober；theworkersofthissamenestexterminatedtwonestsof VsimiHimqtwoof Vanalisandoneof
物〟ぬノ肋γね呼ざ（SeeFig．47）．
Ne＼′ertheless，itiscertainthatA．ceTYlnahasde＼′elopedt＼＼′Ofairlyeffecti＼・edefヒnsivetechniques．neither
possessedbyA．meH脾7％Onebeingpassive−theabserlCeOfso悠tarycounteトattaCks，andtheotheIlaCtive−a
rapidmassattackandaiongcontactwith Vmandarinia．Thisski11fuldefヒnsecontrastssharplywithits
inabilitytoresistpillagebyA，meH蹄7a・Whenco10niesofbothspeciesarekeptinthesameplace，inmost
CaSeSA．me嘩7VultimatelypillagesandexterminatesthecoIonyofA．ce7eTZa（SAXAGAMi1959）．Thus，the
introductionofA．mell紳Ft7tOJapanevokedacurioustriangularrelationamongthethreespeciesduring
a century．
Seriousdamagetobeehivesbyhornets，including Vmanddt壷iamagn抑qarerecordedinIndia（
Cem／Ja血dIca：ShttTH1960），1ndochina（A．c．javana：Touトl＾NO汀1939）andAf岳hanistan（A．c．ssp∴S⊂HNE】DER
andDJÅLAL1939）．KLOFrandSc脚EIDER（1970）describedadefbnsivebehaviourofA．ce7VnaSSP．ineastern
A短hanistarltO掩堺aSP．：Atthearrivalofahornet，30beeworkersfbrm虔densemassatthenestentrance，
eachchangingitsorientationtothehornetaccordingtothcmo、・ementOfthelatter．Ontheapproachof
thehornetbeyondacriticaldistance，beesraisetheirmetasomasandlowertheirheads，tOgetherwiththe
PrOductionofshimrnerlng．Thehornetoniycatchesbeesleavingthedeftnsivemass．Ingeneraltheobserva−
tioncitedcorrespondstomyobservationbuttheinitialretreatandmasscounter−attaCkIobservedarenot
mentioned．Consideringthisabsenceofitsdescription，tO8etherwiththeseriousdamagetoA．ce7Vtlaby
挽軍ainthesamearea，thedefbnsivebehaviourintheAf帥anpopulationseemslessperftctthanirlJapan．
1tisimportanttomakefurthercomparativeobservationsonthedefensivebeha、［iourofA．ceTt7nainvarious
areaS．
IsHAYetal．（1967）refertothemassdefbnsemadebythelocaibeeraceinIsrael：”Thelocalbeesprotect
themselvesagainsthornetsbyfbrmingdenseclurnpsattheentranceofthehive．Insuchin5tanCe，ahornet，
inordertocaptureabee，rnuStfirstdetachitfromtherest，aratherdifncultfbatwhichisrarelysuccessful．
Thereisanotherdangerthattheoverlyventuresomehornet，incomlngtOOCIosetotheaplanphalaru，may
OCCaSiona11ybecomethevictimofitspreyandbestungtodeathbythemassedbees：’Itisnoteworthythat
amassdefense，quitecomparabletothatofA．ceIV／JaCe，tlItqisthusconfirmedinaraeeofA．171e／／昨IVI
Thesimilaritymaybeexplainedeitherbytheparallelacquisitionofasimilardefぬsivemechanismorby
thelossofsuchinsomemembersofthesubgenusApis，aSlha＼′ethusfarnoinformaliononA．dorsa
A．j707eaandtheirrelatives，i．e．，thesubgenera物叩isand肋J叩i51Theshimmeringseemstoofftranother
68
Makoto MATSUURA
ins【anceOfthepresenceofadiscre【ebeha＼バouraJcharaeterinraeesofboth5peCies，forBし】Tl．L三k（1954）
recordedatraceofthishabitin A．me〟昨fVqyPricain Cyprus．Furthercomparativeobservationson
behaviourbothofhornetsandhoneybeesmaythrow王ightonreciprocalevoltltior10fattackanddefbnse
mechanismsin theseinsects．
Judgingfromtheseobservationsseveralpossibilitiesexisttoexplainthediscrepancybctweentheserecords
andobservationsinJapan：1）LowerdefbnsiveabilityinsouthernpopulationsofA．ceJⅥnaagainstレ軸a．
2）Presenccorsomee汀icienthunlingbehaviourinthelセSPaSpeCiescited．3）Presenccofsomepartieular
Si！uation5（hatねci】itatelheattack，OCCurrinseitherecologically（SeaSOnalcorrespondenccofcolonycyeles
Ofpreyandpredator，etC．）orartificia蔓1y（COnditioncreatedbyapiculturalprocedures，etC．）．Furthercom−
parativeobservationsinぬeseareasare陀quired．重吉mightbelyOrthstudyingtわeuseofappropriatebives，
especiallythosewiththeentrancenottoowide．
4．1ntra−atldinteトSPeCific rclationships at food sources
VariousspcciesoFhorne【SObtaincarbohydrates from treesap．A！（helreeaPertUre5＼＼・herethesecretion
isabundant，theyasserttheirterritorialrightbydrivingawayvisitorsofthesameordifferentspeciesfrom
theplace．‡ftwoormoreindividualsofthecongenericordiffbrentspeciesarefbundatoraroundthesame
place，Cer【ain rules arerecognized amongthem．
Thissectiondescribesthedominancebehaviourinhornetwhichisanalogoustothe“peckorder”found
in thedomestjc fbwl（SぐHJEl＿Dt：RUP−EIIL）E1922）．
1）Dominancerelationshipsamongqueensvisitingtreesap
Thcpost−hibernatingqueenssearchfortreesapbefbrede、′Oting【hemseh・eStO（henes【ingsites．ThbJe21
Showsthefrequencyofvisitandthefeedingtimesof18markedindividualsoffour掩軍aSpeCiesthat
VisitedatreeapertureduringthreedaysatthebeginnmgofMayin1967．andinFig．49isglVenOne
exampleconcernmgtheactivitiesoftreesap主ntakeobservedamong12queensand2stylopizedworkers
Oflイma17dariniL7．2queensoflくcIVbroandeachonequeenorl・てaJ7aIisandレ：siITJiI／imaonMay5th、
rrhble2l．Lcngthoftime（inmin．）spenttakingtreesapbyeachindividualvis…tingthetreeaperturedur重ngthrec
da），Sat thcbeginningorMaylstin1967．
Species and
indi＼′idua】
Tin−C
ror
each
fecding tinlC rlOnE壱eding
A、′erage
ハリ
O
2
3
5
7
O
5
QU
3
1
5
3 5
O ︵U
O O
2 2 5 n7 つJ 3
0 0 〇
ハU O
2
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3 ′nV 凸7 n7 つJ
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r r n a
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8
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Frequcncy ，rbtallime
ror
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トO4・07，02・
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marked●
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＊mal−12：queCnOrI′：／冊IJdari17iq，IllaXl−2：Stylopised、＼・OrkeroH・て川a／IdariT血．crト2：quCCnOt■l；：ぐrL7bro．
8 つん
an：queenOr Hα棚梅Xa：queenOr H血乃f〟拍紙
3 0
ComparativebioIogyortheriveJapanesespeciesortllegenuS陀軍α（Hymenoptera，Ⅶspidae）
69
mmm
mm
mmmm
S山岬じ山乱S
m
2−4・592コJ67只Y−人∪−ク︼−2・−−−
m
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qu滅∝≦ふJく ⊃ Q − ＞岩空
mmmC
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C8二木
Fig・49・Ac【ivitiesortreesapintakeobservedamo喝i8markedindividualsor4speeiesI々軍〝thatvisiteda
treeapertureinthedaytimeorMay5th，1967．
Blackrectanglesindicatetheoccupationofthetreeaperture．Spottedrectanglesindicatestayingaround
thetrceaperturewithoutfteding，Arrowsindicateflyingawaysoonafternyingaroundthctreeaperture．
1967atthesameplace．Thel■Cisasignificantdifferencehlthefeedingtimeofeachindividualanditis
Certainthateachqueenoccupiesthetreeapcrtureaccordingtotherelationoftheorderamongthem．
Inthedaytimethetreeapertureexudingsapwasalwaysoccupiedbyaslnglequeen，SurrOundedbyothers
thatwereunabletotakethetreesap．TheyweredrivenbackwithoutanycounterMattaCk，bythethreaten−
ingoftheoccupantwithmandibleswidelyopened，Whenevertheytriedtoapproachtheaperture．
托edingilldividualschangedplaceswhentherirstbecameru11yengorgedandrlewaway，OrWhenshe
WaSdrivenofffromthetreeaperturebyasuperiorindividualarrivingfromanotherplace（F呈g．49）．When
thesameoranotherspeciesvisitedatreeaperturealreadyoccupiedbyafeedingindividual，theresponse
Of【hevisitorlOtheoccupantwasoneortherollo≠・ing：A．Alightingonthepositionatadistanceof
5−6cmfromtheoccupant，afterflyingaroundthetreeaperture，B．Flyingawaysoonafterflyingaround
thetreeaperture，C．Aseverecombatwithmandiblesandstingsbetweenbothindividualssoona氏eralighting
Onnearthetreeaperture；thevictorbeingstotaketreesapandthedefeated，Onthecontrary，fliesaway，
D．Animmediatechangeoverfromtheoccupanttothevisitorwithoutanycombat；theoccupantgives
up the positiolland retreats or flies orr．
Judgingfromtherelationoftheorderofindividualsreferredtolater，thedominancerelationsbetween
Visitorandoccupantintheabovecasesareasfouows：BothinAandBthevisitorisinferiortotheoccu−
Pant，andtheirorderisrelativelyclosetoeachotherinBwhilenotverycloseinA；Onlytwoexamples
OfCwereobservedin1967，anditisconceiYabiethattheyareofthesamerank；Onthecontrary，inD
tlle＼′isitoral＼＼・ayS hasa higherrank than thcoccupant．
Theinferiorandsuperiortypesofbehaviourwereobservednotonlyamongoccupantsandnon−OCCupantS
butamongnon−OeCupantSSurrOundingthetreeaperture．Theinfbriorindividualsretreated、＼巾enperce＼・・−
ingtheapproachofasuperioronewhichfrequentlythreatenedwithopenedmandibles，andsoeachin・
dividualtookupapositionawayfromtheothers，fbrminganapproximatelyuniformspatialdistribution
aroundtheaperture．InTabie22areindicatedthefrequenciesofdominanceandsubordinationamong
markedindividuals，bothintra・andinterspecifically，inmutualattacksandrrhble23showsthefrequency
Ofretreatingbeha＼′iouramong them．
OnthebasisoftheseobservationsglVeninFig．49，and′払bles21，22，23，itiscertainthatthereisa
distinctlineardominancerelationshipamor唱thequeensor物αSpeCiesvisitingtreesap．Theぬ1lowlng
MakoモO MAT5UURA
70
’rhbleヱ2．Frequenciesordominanceandsubordinatingrelationsamongmarkcdindi＼・idualsofrh！e Lr如paspecies
（Cf．’払ble21）visitingtreesapinthedaytimeduringMay3rd・8th，1967．
hldj＼・jdl血
marked ma12ma】ma5m且4ma9ma2−Ⅵa3ma6mallma7ma8ma10m弧Imax2erlぐr2 an】rrequencyor
084637＝437555810∼633659
C
O
O
O
O
O
O一〇1
11
1
・000000000000030 岬
00000000000000 1一〇
〇〇〇〇〇〇O
1・〇一〇
0000・0・1〇一〇〇ユ
000000000
0
O
0
O
O
00
O
0
〇〇〇1−0
l一〇〇〇〇・000
〇一〇
O
1
0000000000・一10000
O
0
00・・00000〇一・・・000
・0
P
nla】O
maX
Tblal
dominance
8
†
0
8
0
6
0
〇
8 31125 3 5 2
0 1 2 − ︰ ‘ 0 0 つ︰ U
6
1
O O
〇
a
4
〇
〇〇〇
Z 3
a a
Qノ
〇〇〇〇〇〇〇〇
a a
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a a
O O O
1 55 4 0 王
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〇〇〇 1 0 0・−一U O O
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O・・00000〇一・・・000
0ニュ0 11一〇〇〇〇〇〇〇〇〇〇
〇〇1−1一〇〇〇〇〇〇〇〇〇〇〇
20−3一〇〇〇〇〇〇〇〇〇〇〇〇
202一〇〇〇〇〇〇♯ 0000〇二〇
〇
5
m m m m
2502314一102000000
200011212一〇〇〇1−000
・000 000つー〇一〇〇10000
O
〇〇〇
・∩∨nVn
〇〇〇〇〇〇34030005一ハ∨・−
00001000000000〇一〇
dominance
〇1一〇〇〇−000・000000
〇一〇
a
● No chance or encouIlter．
000000〇一1831223002
max l
一〇〇〇〇〇〇〇〇＊ 000＊ 00・
1
of retteat
00001〇一71100004 11
0000〇一11拍114010810
O
● No⊂hanc亡Or enCOunter．
000〇一〇24700010000
04〇一925680・100400
0・
a
Tbtal
0之一153ほ801−000810
7・∼−1・1
2
汁equency O O ヱ
・000
rrequencyor 53 72 41 39 王4 37 ま6 31 】 3 0 j 4 4 1 i 3
〇一っー01847詑13100004I
ヰ︰617
a a a a 8 a a a
m m m m m m m m m
lユーー5
a
ma 8
maほma王 ma5 ma4 ma9ma2 ma3 m往るm盆】】ma7 ma8ma10max】乃1鉱2 cr】cr2 anl加quen野Of
7btal
i爪dividual
marked
Subordinati】唱
Table23．Frequenciesofretreatingbehaviouramongqueensandstylopizeくiworkersor物αSpeCie5（Cr，Tもble21）
Visi【ingけeesapdurj喝May3rd−8tね，iタ67．
ComparativebiologyorthefiveJapanesespeciesofthegenus7々spa（HymenOptera，Vcspidae）
71
Ordercouldbeobservedasanintraspecificdominancerelationshipamongthequeensandthestylopized
WOrkersof V：mandariniawhovisitedthetreeapertureatthebeginnlngOfMayin1967：
ma12−（㌔ア1ト9−ma2−ma3−ma6窪……1＿mal。トxi−maX2
1nterspecificdominancereiationshipamongqueensvisitingtreesaplS，thoughgreatlyinfluencedbythe
individualdisposition，generauyaSぬ拉ows：沼α〃血rf〃ぬ−Cm∂和一α〃α／ね一頭涌肋Ⅷ㌧礪画飽
Eachindividualtakestrcesapaccordingtotheordermentionedabove．Thequeenhornetma120f
manddriniawiththetoprankcouldoccupythetreeapertureandtakeherfilloftreesapatanytimewhen
ShevisitedonbothMay6thand8th．Thenextpositionwasoccupiedbythreeindividuals，mal，ma4
andma5，allVmandariniaamongwhichaninterestingtriangularorderwasnoticed．Eachofthesethree
queenscouldtaketreesapintheabsenceofsuperiorones，butwouldflyoffwhenasuperioronearrived．
王ndividualsundertheabovefburqueensintheirorderusuallyspentmuchtimenearthetreeaperture
Waitingforanopportunitywhentheoccupantwouldtakeherfilloftreesapandflyoff．Therelation
Oforderamongfourqueensatthebottom，mall，malO，ma7andma8，WaSnOtClear；becausethese
individualshadlittleopportunityoftakingtreesapinspiteforwaitingpatientlyduringthedaytime（rrable
21andFig・49）・However，eVenindividualsatthebottomcouldtaketheirfil10ftreesapintheevening
as mentioned belo＼＼・．
2）Behavioursoroverwinteredqueensintheevening
Thehibernatedqueenstaketreesapnotonlyduringthedaytimebutalsointheeveningfbrseveraldays
SOOnafteremergingfromtheirhibernacula．In1966Que7TuStreeSbegantosecretetreesaponApri125th
andhibernatedqueensof物aspeciesvisitedthetreeaperturewherethesecretionwasabundant．Their
activityoftreesapintakeintheevenlngCOntinueduntilMay3rd．Thereafternotasinglequeencould
beobservedatthetreeapertureintheevenlnglnSPiteofabundantsecretion．Oneevenlngthetreeapeト
turewasoccupiedbyaqueen，andwithinaboutlOcmfromtheapertureseveralqueensofthesameor
differentspecieswaitedforanopportunitytotakethesapuntiltheoccupantleft．IntheevenlnganOCCu−
Pantisdominantovertheothersandisabletotaketreesapfullywithoutdisturbancebysuperiorin−
dividuais，Sincenootherqueensof物aspeciesarrivefromotherplacesintheevening．Whentheothers
approachedthetreeapertureshethreatenedwithhermandiblesanddrovethemawayasobservedinthe
daytime．Thedominancerelationshipamongtheoecupantandnon−OCCupantSandamongnon−OCCupantS
WaSaIsoobservedduri喝theevenlng．職ble24indicatestbe丘equencyordominancebehaviouramo喝
Table24．Frequenciesofdominanceandsubordinatingrelationsamongqueensof陀3Pamandarinjaand V
CrUbrovisilinglreeSaPinlIleeVeningon Ma）・1st，1966．
IlI
ml
0 0
〇一
m］−5；queCn Ofl′：lTlalIdarinia．crl：queCn OrトてcrL7bro
9
0 0
〇一2
20
crl
0 0
32
0
0 0
4
0 0
m2
cr 1
m2
0
1
m3
0 〇一柑ヱ
m 5
m3
m5
〇一日 15
一1 2
ml
m4
Tbtal汁equency
ofdominarlCe
m4
72
Makoto MATSUURÅ
fivequeensofレ：mandariJ血andaqueenoflてcTt7broduringthee＼・eningofMa〉・1stin1966at agroYe
inKibi．Itisapparentthatthereisalineardominancerelationshipamongthesequeens．Theorderwas
as follo＼＼′S：ml−m4−m5−m3−m2−Crl．Asshownin’rhble24，aCCOrdingto theordergiven abovethe
individualwhichisnexttotheordercouldftedsuccessiveiyiandsofbrth，Whenanoccupanthadle托the
treeaperture．OnMaylsttheomegaindividualof VmGndarinEq，m2，finishedtreesapintakeat10：25
PLm．and（henaqueenof Vcrpbl’qCrl，COuldfina11yoceup〉■thetreeaperturesuccessi＼・elytil110：50p．m‥
Onthefbllowingevening，May2nd，Iobservedfi、・equeenSOfl（：ma17dal・i11iaatthesameplace．orwhich
three，m5，m3andm2，hadbeenobservedpreviousevenlngandheldthesameranksamongthemseives．
Intheevening，aqueenOf Vmandariniausuallycontinuedtotaketreesapfbraboutanhour．Then，With
a魚11lcrop，Sheleftthetreeaperture，Walkedtothetopofthetreeandsettledontheundersideofabranch
7−8mhigh．Cleaningherself【here，ShefdJas】eepconceaJingthe【ipsofhcrFo】dedwingsunderlheventral
Surfaceofherabdomen，Similarto theposition fbundin hibernacula（M＾TSUUR＾1966）．
Itisuncertainonwhatfhctorsintra−andinterspecificdominancerelationshipamongqueensof Lb5Pa
SpeCiesisdependent．PARDI（1948）explainedbioIogicaliythatthedominanceorderamongfoundressesof
jblis（esga〟jcuswasdeterminedbythedegreeofovariandevelopment，andYosHIKAWAetal．（1969）reported
thattheconditions ofthe reproductivesystemgenera11ycorresponded to the socialorderinanestof
撤rischnogαざterSP”Inthecaseof挽汐aqueenSVisitingtreesfbrsapitisdifficulttofindacorrelation
betweensocialrankandovariandevelopmentbecausetheovariesofqueensaremostlyundevelopedatthe
incipientstageofdeveloprnentduringthefeedingperiodsoonafterhibernation．Itissuggestedthatthe
dominanceinteractionamong陀甲aqueenS，eSpeCiauyin Vmandariniq，reSultsinaquantitativedifftrences
Offbodintakeamongthem，andthismaydividetheposトhibernatingqueensintomoreorJessdistinctnu【ri−
tionaigroups：afbwhighnutritionqueensandtheotherlownutritionones．Itseemsprobablethatsuch
rlutrはonaldiffbrencesservetodiffbrentiatethe”quality”ofqueens，anditispointedoutthatthereexists
asignificantrelationshipbet＼Veenthequalityofqueenandtheeventualsizeofthecolonyintheearl〉・Stage
OfnestinginDo／ic／脚融pLL／ov，／vestris（BRIANandBRIAN1948，1952）andinlセ5Pu／avuなarLs（ARCHER1980）．
3）Interspecificrelationshipsamongworkersatfbodsources
lnterspecificdominancerelationshipsamong悔paspeciesisrecognizednotonlyamongqueensbutamong
WOrkers．ThefbJ】owingorderhasbeenobserl℃dasan血erspec漬cdornir）anCerelationshipamongfj＼′eSpeCies
O＝頻卯WOrkersnearKibi：mandari／血−ana［is−Cmbro−Simi（／imo−（rPPico，Whichisthesameasthat
fbundamongthequeens．Therelationsarnongallspecies，eXCeptfbr Vmanddriniq，areSOmetirnesindistinct
ineachencounterandmayresultinaseverecombatwithmandiblesandstings．
Vma［7dariniainvariablyoccupiesthefirstrank，mOnOPOlizesfhvourablesapsources，andexpelsother
hornet species，beetIes，butternies，etC．Zfatreesapsi【eis oecupied byIて／T）G／t（わrin庇＼、’Orkers ofother
SpeCiesnyawaywithlittlehesitationsoona氏erflyingaroundthetreeaperture．Ontheotherhand，VtrQPica
isthemi1destspeciesandisoftendrivena＼Va）・byothel■SPeCiesFromafbodsource．1nspiteoftheirmuch
largerbodysize，Vlrvpicaqueensarealwaysexpe11edbyasingle Vmandarinidworker．
A11theworkersforagingatafoodsourccalwaysbelongtothesamecolonyalthoughtheycannotcom−
municatethepositionofasourceoftreesaptotheirnestmates．勒rkers鉦ornthesamecoIonydonot
behavea鎚reSSivelytowardseachotheratafbodsource（Fig．50）．Otherhornetsofthesamespeciesbelong−
jng【Oaliennes【Sareehased away by’’1eBi【jmate”ownersⅥ雨h orwithoutcombatwheneverthey tryto
approachthetreeaperture．Thedefenceoffavourabletreesapsourcesisramarkablein t／：mar7dGri17ia．The
OCCupiedfbodsourcelSguardedbysomeindividualssittingaroundthetreeapertureswhichdonotreturn
to theirown nest．Thesebehavioursindicatea＼Ve11de＼′eloped territorialdefene
Anotherpeculiarhabitatfoodsourcesisthefrequentextranidalfbodregurgitationobservedbetween
F！1TIa／ldarinJbworkers（OK＾D＾196l，M＾TSUUR＾1975）．Mostl′eSPine＼VaSpS arelVe】1kno＼Vn【O tranSftr
regurgitatedfbodamongworkers（Montagner1966，Spradbery1973，Greeneetal．1976），buttheygenerally
donotinteractwithotherfbragmgworkersatmostnaturalfbodsources．Fbodregurgitationbetweenworkers
isconfirmedwithinnestsofaufive物aspecies，butoutsidethenest．Ihaveseenitoccumngonlyonce
Withtwoworkersof Vm7browhenvisitingatreesapsourcein1974．In Vmandariniathesituationis
Comparativcbio王ogyofthefiveJapanesespeciesofthegenusI毎a（Hymenoptera，Vbspidae）
73
Fig・50・Lespamaltdari77iaworkersco11cctingsnpexuding fronlthebarkofanoak，Q，lelrEJ∫Serm（a．
Fig．51．TtophaLlaxis betweenl＼＼，OStar＼，ed＼＼・Orkersor［bpalTlalldari（1iLlOnthcgroしl【ld．
mostfrequcntI〉，enCOunteredamongnestmatcsvisitingthesametreesapsources，in＼＼・hichcasetherood
flowsunilaterally，nOtreCIPrOCallyThemoreinterestinglnStanCeisseenbctweentwostarvedworkers：em−
bracingcachothertightlyandeitherlyingorrouingonthegroundorhangingfromatreebranchbya
Singlehindlegofoneindividual，theyaiternatelybegandreceivefbod（Fig．51）．Thecmbracelastsfrom
SeVeraltotensofminutesandtheydonoteasilyseparatecvenifdisturbed．Suchreciprocaltrophaはaxis
isoccasionallyseenwhenthisspeciesattacksbeehivesornestsofothersociaiwasps，eSPeCiallynearthe
Victim’snestsduringpro10ngedattacks，butsofhrisnotconfirmedwithinornearitsownnestsnorin
74
Makoto MATSUURÅ
queensand males．
Foodregurgitationarnongnestmatesisapparentlyahabitindependentlyacquiredbyvariousgroupsof
SOCialinsects．Theintenseextranidaltropha11axisinレ：mandari17iaiscertainlyaspccializedcharacter．It
mustbementionedthatthisphenomenonismainlyobservedatsapsourcesandnearvictim，scolonies，
thatis，ateOnCentratedfbodsources．Inhoneybees，eXtranidaltrophallaxisappearsatartificialfeedersbut
Seldornatnowers．Thehabitin Vmandariniamayberelatedtoitspreftrencefbrsuchconcentratedfbod
SOllrCeS．
5．Discussion
Asisusualinothersocialin5eCtS，VeSplneWaSpShaveevoIvedtobeeitherhighlyspecializedorverygeneralized
intheirfbodprefbrence．Incontrastto掩叩頭如pecieswiththeirsmallbodysizeandomnivoroushabit，掩軍a
SPeCie＄haveanincreasedbodysiヱeandpredaceoushabit，andhaveattainedaco−eXistenceintheirfeeding
relationshipsby signiricantdifftrencesamongthem．
VtTqPZCaisatyplCalspeciaiist，beingaimostexclusiveiydependentonthebroodofpoustinewaspsfbrits
protein food，andthelifecycleofthisspeciesis synchronized with thatofpolistine＼VaSpS．
Ontheotherhand，V eTqbTDisregardedasasemispeCialist，PreftrringvariouskindsofcicadasinJapan．
Itseems Zikel）・【hat this species js hjghlyspecia厄ed behavjouralJ）・fbreapturjnganycjcadas，and【heyhal′e
anabilitytoutilize Cl：岬Joり・l叩anajbcia／jLWhichisabouttwotiIneSlargerinbodysizethanI′：cIVbrqand
huntingthislargestcicadahasso farbeenobser＼・edonlyby t／：cIVbro．Thehornets also havetheabilityto
huntawidevarietyofinsectswhentheprefbrredfoodsourceisIowerindensity．Rearingofsexualsinthis
SpeCiesceasesinearlySeptembeちinpara艮elwiththedisappearanceofcicadasinsouthwestJapan．Sucha
fbedingstrategymakesbetteruseoffavourablefbodandcontributesgreatlytoreducethecompetitionfbrthe
food resources among職印SpeCies．
Both Vana胎and VsEmiHimahavegeneralizedtheirftedingconsiderably．rtisplausiblethattheseswift
andagilespecieshavethecapacitytofbrageonvariousfbodsourcesscatteredinthefieldsandthesearefbund
inlargenumbersintheregionswheretheybuildtheirnests．Thecontrastingftaturesofthet＼＼′OSpeCiesshow
that VanatisissmallincoIonysIZeWhile Vsimi［Hmaismuchlargerthananyotherl句paspecies，andtheir
peakpopulationsoccuratdiffbrenttimesoftheyear；thoughbothspecieshavelong−CyClecharacteristics．These
differencesmayenablethemtoreducecompetition forsimilarfoodresources．
Ontheotherhand，Vmanduiniahasachievedanef蔦cientfbraglngStrategyinthattheyattackinmass
thecoloniesofothersocialwaspsandhoneybees，andalsopossesstheabilitytoh11ntaWidevarietyoflarge
insectsandspiders．Apresumed courseofevolutjon ofthispeculiarhabitis briefly outlined below．
Thelargebodysizeandenormouslydevelopedgenae（MATSUURAandSAKAGÅMI1973），gaVe Vmandarinia
theformidabilitytoattackandtoconquerotheranirnals．Bythisability，COmbinedwithanextraordinaryag−
gressivenessandthestrongtoxicactionofthestout＄ting，VmanddriniaoccupleSanunrivalledpositionamong
insectsandothersmaJlanimals，atleastinJapan．Incompensation，however，itsrelati、・e】）rS】uggjshmovement
makesthecaptureoffugitivepreydifficultwhichisespeciallyconsplCuOuSincomparisonwithsomeagilespecies
SuCh asl・：simi［［i［77aand V：ana／is．Consequentlyitisplausiblethat aslightdisplacementoftheecoLogical
nicheshasappearedbetⅥ・eenlてmandariniaandothercongenericspecies．Vmandariniadidnotchoosethe
SpeCializedwaytooligophagyaspracticedby Vtrqpica，Ⅹeepingitspo量yphagoushabit，Vmandarin由began
toseekpreywhichwaseasytocatch．Amongsuchpreytherewouldbethosemakingcolonies，havingalarge
amountofconcentratedproteinfbod．HoweverthedependenceonPo［istesasin Vtrqpk，aWaSprObablyim−
possibleぬr H椚α乃（おr如∼ちWhosenestsreachalarge＄izeandrequlrea払喝equantityofanimaldiet．Onthe
Otherhand，defヒnsebynestownerspresentednoseriousobstaclefbr Vmandat血iqanditwasratherfhvourable
fbr Vmanda71niaifplentyofsuchselfidefbndingindividualswereavailable．Thehabitofattackingnestsof
Othersocialwaspsandbees＼＼・Ouldbeaconsequenceofthistendency．Thus，V川andar（niaaffecISOthereon−
genericspeciesintwoways：aSaCOmpetitorinseekingcommonfbodsourcesandasapredatonItisinteresting
thatitdoesnothuntotherspeciesattreesapso11rCeS，merelyexpellingthernasIowerrankedcompetitors．Aト
tack on nests ofotherspecies，firststarted asindividualhunting，WOuldoccasionallyberewardedbyan
ComparativebiologyofthefiveJapanesespeciesofthegenus掩鱒a（Hymenoptera，Ⅵspidae）
75
abundantsupplyofimmaturestobe fbragedwhentheattack＼l・aSCOntinuedtoovercometheresistanceof
Victims，eVenifaccompaniedbyalossofseveraltoseveraltensofnestmates，Whichweretransfbrmedtomeat
Pelletsandfbdbacktothecolony．Theslaughterphase，themostinterestingstageintheattackby Vman−
dariniaonothersocialinsects，muStbealaterinsertionbetweenhuntingandoccupationphases．Thisphase
isabsentwhenattackingjb触Jesnestsandseernlnglynotwe11developedwhenattacking挽甲uぬnests，but
itisdistinctwhenattacksaremadeonvictimswithseverecounter−attaCks，i・e．OtherhornetspeciesandApis
mel／紳taItsreleasingmechanismisstillunknownbutcertainiyisrelatedtoachangeofmotivation，Which
isprobablyreleasedbytheintensecoumteトattaCksandthepresenceornestmates．Itsfunctionalsigniricance
maybeto払cilitatetheoccupationofavictim，snest．
Thedeftnseofoccupiednestsagainsthornetinvaders，reCOrdedalsoin Vo7鹿ntaiis（IsHAYetal．1967），is，
inalesserdegree，reCOgnizedinthecaseofoccupationoffhvourabletreesapsources，tOO．Suchpropertydefbnse
isobservedalsoincleptobioticstinglessbeesofthegenusLesirimeHtta（NoGUEIRA−NETO1970）．Comparedwith
theattackandnestoccupationbythisobugaterobberbee，theattackby Vmandarinjaisfarlesselaborate
ander鮎ient・Fromthis払ct，tOget鮎r雨tbthemaintenanceorapoiyphagouspredaceoushabitingeneral，
theattackof Vmandariniaonnestsofothersocialwasps，includingboth掩spaandI句ダuれisinterpreted
asaretBtrdedspecialization．Uptothepresent，effectivedefbnseagainst Vmandariniaisknownonlyin
CeTUna．Thismaybeanadaptationacquiredthroughalongcontactwiththepowerfulenerny．
TheintroductionoftheEuropeanhoneybee，Apisme〟昨TqintoJapanin1876andsubsequentdevelopment
OfmodernaplCultureo約陀dagoldenopportunityto㌣研α〃血r如ね．Thestandardapiculturalracesor．4．
melJ紳772WereCOn丘ontedinJapanbyamonstrousenemy，tWOtimeslargerandincomparablymorefbrocious
than VctYlbTVintheirhomeland．SolitarycounteトattaCksbybees，beingeffectiveformostenemies，SerVed
merelytoincreasethefbraglngefriciencyoftheirenemy恥㌻巨椚α〃ぬ〃〃′ちtheEuropeanhoneybeewasceト
tainlyanidealpreybecauseofitslargecolonysizeandftequentcounter−attaCksasweuasthefbeblenessof
eachindividual．Theoutbreakofcatastrophicdamageby Vmandarinjaistherefbreregardedasararecoin−
Cidenceofethologicalcharacterspossessedb〉lthepreyandsoughtbythepredator．
ⅠⅤ．DIVISION OF LAおOUR
Allthespeciesofthegenus物ainJapanfbundtheircolonieswithaslngleftrtilizedqueen．Thissolitary
Stagelastsfbrabouttwomonthsfromlatesprmgtoearlysummeruntiltheemergenceofthefirstworker．
WitbtbeappearanceofworkersthelabouroftheqtleenVariesinbothquantityandqualityinthecourseor
COlonydevelopment．Fina11y，ShebecomesrestrictedtoovlpOSition，leavlngnearlyalltheotherdutiestoworkers．
Thus，theworkerstakechargeofallthetasksin［hecolonyexceptegglaying．Ho＼VeVer，＼Vhenthequeenis
10Stbyaccidentornaturaldeathinthccourseofcolonydevelopme叫OneOrmOreWOrkersbegintolayeggs
andproducesomemales（D＾RCH￡N1964，M＾TSUURA1968）．hthischapterthedivisionoflabourwi11bediscussed
Onthreeaspects：behaviouraldiffbrencesbetweenqueensandworkers，thoseduringtheadultlifbtimeofthe
、、′Orkers，and thoseamongworkersin orphancolonies．
l．Behaviouralchangeofthequeeninearlynestingstageinrelationtoworkeremergence．
Theactivitiesoftheqtleeninsolitarynestfbundationaretypica10fasolitarywasp．Afterworkeremergence，
thecolonyundergoesaslgni翫antactivitychangetofbrmthedivisionoflabourbetweenqueenandworkers．
Thefollowinginfbrmationismainlybasedonthedata鉦omanestof Vc7Vb7V（C6601），Whichwascon−
tinuouslyobserved丘ornthesolitaryperiodtothecooperativeperiodin1966．Variationsbothinqualityand
inquantityofqueenactivitiesof Vm7b柑areShowninl胤bles25，26and27．
1）Extranidalacti＼′ities
a）Collectionoffood；Fbod−CO11ectionactivityconsistsofaunkedsequenceofsearching，hunting，treat−
mentofprey（＝Primarymalaxation）andtransportinthecaseofproteinfbod；Ofsearching，SuCkingand
transportofcarbohydratefbod．Thenumberoftripsobservedduringthesolitaryperiodof VcTUbrvwas
39，Ofwhich30tripswerefbragingforfbodandthetimedevotedoccupied46．0勒onJune3rd，1966and
Makoto MATSUUR入
76
lbblc25．Changeinrelati＼・etimespenlinvariousaclivitiesbythcqueenor V＆spacmbroduringtheobser＼・a・
tion pcriod．
Date
Ob5erVation
time
（1966）
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Intranidalactivity
Oviposition
（min．）
0
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Pcrcent oreach timcspcntin
Total time of
observation
0
q′
0
5
2
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is5−16minutes
55．9％onJune5thofthetotalobservationtime．Durationofatripforcarbohydrates
5
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q′
queenoccured
10th，＼＼血nthefirst＼＼・Orkersappeared．Thclastobser、・ationorforagingfor foodbythe
OnJune26thwhenonlyfiveworkerswerepresent．DuringtheperiodfromJune10thto 26th aitogether
Thetimespent
37tripswereobserved，Ofwhich36werefbrcarbohydratesandonlyoneforinsectprey．
minutesわrin−
forforagingwas2r45minutes何＝18．2，S．d．よ10．4）fbratripfbrcarbohydratearld27．5
SOlit8ryperiod
sectprey．Theintertriptimespentinthenestwas2−8minutes（菰＝4．3，S．d．j＝2．0）inthe
and7−26minutes挿＝12．3，S．d．土14．1）a魚ertheemerge11CeOrWOrkers．
Thetirnefbrgatheringnestmaterialsoccupied14．0％ofthetotaltimeofobservationonJune3rdand
9．4甲0OnJune5th，reSPeCtively，inthesolitaryperiod．Withtheappearanceofworkersthequeengathered
veryfewnestmateria重sfromthefield（職ble26）．Theiastco11ectionofnestmaterialswasobservedon
Ju重y3rd，thefinaldayofherfieldwork．Thus，COueCtionofnestmaterialsbythequcencontinuedfbr
aboutthreeweeks紙eremergenceofworkers．
2）Intranidaiactivities
Theproportionofinsidetaskstooutsideonesbythequeenincreasedwithworkeremergence：OnJune
3rdand5thjustbeforetheemergenceofworkersitbeing40．0呼band34．2勒，reSpeCtively，Oftheobserva・
tiontime．OnJunelOthattheappearanccoffirstworkers，52．5勒ofhertimewasspentinthenestthough
一一〇
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3
4
0
0000000000
3
3
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0
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O
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〇
O
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〇
0
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0
9
＄
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5
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4
7hble27・Changeofrateofvariousintranidalactivitiesbythequeenof掩軍aC7Vbrq，perhour，duringthe
O
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、i・し︰∴∴∴∴∴
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‖＝‖＝＝‖
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Carlnibalism
（。…e 鞄nning。1慧㌔g 慧r
O
Theratiosinworkersgivcnbyaveragesofindividtlalsobserved．Arrowindicatesthcemergenceoffirstworkers．
9吼5
カ030
鋸．1001000弧010000〇．〇〇〇〇〇〇〇〇〇
♀ 単 ♀ 受 ♀ 萱 ♀ 苧 ♀ 螢
（l11i】1．）
∵∵∵∴
7
00
モノ
J
e
4．20
June3
00
つー
Observatiorlperiod，
WOrker5
Observed） byqueen
ぬm烏111烏
No lcad
Waモモr
IT‡esh
Liquid
騨）（trip5′hour如a如亜5￡…ご霊芸
7blaino．or
Percentaぶe柑tioorva雨usload5broughtbaぐkin
tolallrわsrecordedbolhiれqueenandworkr￡
Aver喝etime
‡匂raging 叩entinthe
Date rateorqueennest between
77
ComparativebiologyofthefiveJapanesespeciesofthegenus掩spa（Hymenoptera，Vespidae）
rrhble26・ChangeoffbraglngaCtivitiesin Vaspacrqbroqueencomparedwiththoseinworkers．
78
Makoto MATSUURA
newlyemergedworkershadnotyetstarted”social”activities．ThequeencontinuedoccasionalintranidaI
activitiesasshedidinthesolitaryperioduntilthelastobservationofJuly27th，thoughshecametospend
rnostofhertimerestirlgOnthecomb（rrhble27），Detailedproceduresfbrvariousactivitiesaredescribed
as fblloIVS．
a）Provisioningoflar＼′ae
Thefbodoflarvaein物aisoftwotypes，liquidandso最d．lmmaturelarvae，lstto3rdinstars，arefもd
Onlyonliquids．Ontheotherhand，4thand5thinstarlarvae，eXCeptfbrthoseofVtTPPtCqareprOvisioned
withbothliquidandsolid．Inthecaseofsolidfbods，aqueen，havlngfinishedmalaxation，apprOaChes
agroupoflarvaecarrylngthefleshpelletwithhermandibles．Fromthefrontofthepelletasmallmorsel
iscutoffandplacedonthelarvalmouthparts．Thisprocedureisrepeateduntiltheentirepeuethasbeen
disposedof．Asinglepelletisdistributedto3∼7maturelarvae．Afterworkeremergenceapartoftheprey
Capturedbythequeenisfrequentlysharedamongworkersas＄00naSShereturnstothenest．Thequeen
mayobtainafbodpelletfromafbraglngWOrkeroramalaxatingoneonthecomb．Oncethequeenbegins
toftedthepellettolarvae，Sheneversharesthepelletwithworkers．Havingfinishedprovisioningthelarvae，
Sheusuallyperfbrrnsaeleaningbehaviourwithherlegsandmandibles．Thenshebeginsagaintoapproach
thelarvaeandtoftedliquid，SuCked鉦omthepreyduringmalaxation，tOthelarvaeinsuchamannerthat
ShegetsthetongueneartothemouthpartsoftheIarva．Inthecaseofliquidfoodcontainingcarbohydrates，
Shefirstcleansthelegsandantennaesoonaftertheforagingtrip．Then，Shebeginstoprovisioninasimilar
Way．ThequeencontinuesprovisiorlingthelarvaewithfoodaftercessationofherforaglngaCtivitythough
thefrequenqbecomesverylow．Thelastdistributionofliquidcarbohydratesbythequeenwasobserved
OnJuly27th，Se＼・enWeeksafterworkeremergence．Malaxationandfoodpro＼′isioningoccupied17・0勒and
19．4％，reSpeCtively，OfhertirneonJune3rdand5thofthesolitaryperiodandO．0∼15．4甲oduringtheperiod
afterworkerenlergenCeineach daytimeobservation．
MoRIMOTO（1960）suggestedthatfbr馳始teschinensistheftedingbehaviourofthequeenisreleasedby
CertainstimuliemittedbylarYae．Inall掩甲aSPeCiesthe5thinstarlarvaescratchthewallsoftheircells
withtheirmandibleswhentheyrequirefbod，andthisperぬrmanceapparentlyservestoattractadultnurses．
In VtTqPtCathequeentapswiththemiかandhindlegsonthecombforawhilewhenshereturnstothe
nestwithfoodinhercrop．Thisisregardedasaslngnalsysteminfeedingactivitybetweenadultsandlarvae．
b）Nestconstruction
Buildingactivitiesinsidethenestincludesuchdiverseoperationsasconvertingrawmaterialsintopulp，ma
thepedicelandpaintingalustroussubstanceonitssurface，makingceu＄andlengtheningtheirwalls，mak−
1ngandenlarglngtheenvelopeandputtingpasteoncocoons，Thesetasksarechieflyaccomplishedwith
hermandiblesandforelegs．Materialsfbrnestbuildingaregatherednotonly鉦omthefieldbutn．omwithin
thenest．Thequeenfrequentlyscrapesthebottompartoftheenvelopeortheupperedgeofacelland
incorporatesthematerialagalnintotheneststructure．Withoutgatheringthematerial，Shesometimesre−
thinsstiilmoistpartsofce11sandenveloperecentlyelaboratedbyworkers．
Thequeenof Vc用b7VSpent，20．6％and12．6勒ofhertirneinenlarglngtheenvelopeinthesolitary
periodonJune3rdand5th，reSPeCtively．Withtheappearanceofworkersthetimespentinnestbuilding，
insuchactivitiesascoatingthepediceいen紳eningceilwallsandenlargementofenvelope，decreasedto
4．6％ofhertimeobservedandcellinitiationwasneverbeenobservedafterworkeremergence．Retirement
frombuiidingactivities advances through fourstepsinwhich the fbllowingprocessesaresuccessively
eliminated．（l）Initiationofcells，（2）Enlargementofceusandenvelopewithnestmaterialswhichsheherself
bascollected餌）mtbe員eld，（3）Enlargementofce11wallswithnestmaterialscollectedbyherselrwitbin
thenest，（4）Re−enlargementofcellwallswithoutnestmaterials．Thisbehaviourcontinuedfbra払irlylong
timeaftertheemergenceofworkers．Iobservedthequeenrethinningarnoistpartofaperipheralcellin
adefferentnestof V m7brvwithabout150workersonAugust15th，1965．
Ina11掩軍aSPeCiesthenestmaterialcollectedbyaqueenorworkerisalwaysprocessedbythecollector
herselfoverawholeprocedure，andisusedfbroneperpose，aSalsoexhibitedin撤Iisteswasps（WEST一
己別謂朗ARD1969）．
Comparativebio量ogyortheriveJapanese5PeCiesorthegenus掩軍α（Hymenoptera，Ⅶspidae）
79
C）Nestpreservation
Thisincludessuchtasksasnestcleanlng，ne＄tdefbnceandregulationofnesttemperature，thesemostly
beingmadelessintenselythanbyworkers．Behavioursfbrnestpreservationbythequeenof VcTUbrvare
rarea托erworkeremergenceasweljasinthesolitaryperiod．Thefbllowlngbehaviourswereobservedonce
ineachease．（1）Coatingthepedicel（June16th），（2）Remo、′alofthefragmentsofthecapofacocoon、Vhich
hadbeencutawaybyanemergingworker（June26th），（3）Ⅰねnning（Julylst），（4）Regurgitatingthewater
Onthecapsorcocoons（July22nd）．Thesedisappearwiththeincreaseinnumberorworkers，eVentually
beingleftentirelytotheworkers．
d）Oviposition
Inall挽甲aSpeCiestheovipositionprocessconsistsofalinkedsequenceof：Pre・fixation（Ce11inspection
Fig．52．Ovipositionbya物αC和む柑que臥
Fig．53．馳g−layi喝WOrkerso‖句卯Ⅶ融閃inanorphancolony．Threeortbemareinspectingthee㍑Sdeposited・
Makoto MÅTSUURA
80
andinsertion），蔦Ⅹation，OViposition，Selトcleaning，abdomenwithdrawalandfbodintake（almostentirely
larvalsecretion）．Thedurationofovipositionfromthefixationtotheextractionofthebodyrangedffom
82”82seconds（市＝136．7，n＝16）rorIイm7blt｝queenS．（Fig．52）．Ontheothcrhand，intlleCaSeOrWOrker
OVlpOSitioninthesamespecies，thelayerstaysatthecellforawhileafterovlpOSitionandcontinuesto
inspecttheeggdeposited（Fig．53）．Theovipositionrateofa V c用bTVqueendidnotreachl．Oeggper
hourthroughtheobservationperiodevenafterworkeremergence．Thenumberofeggslaidperdaywas
O・5eggsinthesolitaryperiodaTldanaver喝eOf4．8eggs（max．14，July24th）わr49daysaftertbeeme曙enCe
Ortherirstworkers．TheshortestpausebetweensuccessiveovlpOSitionswas20n扇nutesonJune25th．On
theotherhand，Whentheworkernumberreachedoveronehundred，thenumberofovipositionsincreased
rapidly，prObablyreachingmorethanseveraltensperday．
e）Resting
Themostremark且blefbatureaftertheemergenceofworkersisthatthequeencomestospendmostofher
timeatrestonthecomb，WithonlyoccasionalcellpatroIsandtrophauaxiswithworkersandlarvae．The
queenof Vc相brospentonly3．0％and2，l勒ofthetotaltimeofobservationrestingonthecombontwo
daysofthesolitaryperiod（7hble25），reSpeCtively．AfterJune10thwhentheworkersemergedshewasfrex
quentlyseenrestingonthecomb，andafterJuly6thwhenshestoppedfbraglngaCtivities，mOStOfhertime，
89．3∼97．9勒，WaSSpentinrestingonthecomb，Howeveちa氏erearlyAugust，therestlngtlmeOrthequeen
SeemStOdecreasetosomeextentwithacorrespondingincreaseinthenurnberofceusreadyfbregglaying
as the number orworkersincreases．
2．Relationsbetweenlabourandagein workers
l）Eme柑enCeOfworkers
Workersemergcfromthepupainsidethecocoonandspend6∼10hoursintheircellsinorderthatthe
WlngSandintegumentcanhardenbefbretheyemergen’Omtheceus．Atthetimeofemergencetheysecrete
Salivary伽idwhichhelpstomoisten血ecellcapandtomakeegresseasieこThen，theygnawぬecocoon
CaPWiththeirmandiblesf（Omtbeinsideandcutaraggedholetowardonesideoftheendofthecocoon■
批takesabout30司Ominutes丘omthebeginnlngOfcocoongnawingtothetimewhentheygetout鉦om
tl】CCell．Duringthe＼＼巾oIcprocesstheygetnohelpfromthcolder、、JOrkersinthcnest．Ontheother）lan〔l，
WhenBombusspeciesworkersemerge，theyarehelpedoutoftheircocoonsbyseveralworkersinthenest
（KATAYAMA1966）・
2）Behaviours ofworkerssoona托el−emergenCe
Newlyemergedworkersimmediatelyseekoutfullgrownlarvaeandreceive5alivarysecretionfortheirfirst
fbodasanadult．Theydonottrimtheraggededgesoftheccl重whichtheyhavejustle托butleavethis
taskfortheolderworkers．Theymoveverylittleandspendmostoftheirtimethrustingtheirheadand
abdomenintoemptyoreggcellsatthecentreofthecombexceptforoccasionalfbodintakefromlarval
SeCretion，Selflcleanlngandslowwaikingonthecombforthefirstdayoftheiradultlife．Thisapp呈iesin
allfi＼・elセspaspecies（Thbles28，29，Figs．54−56）
Intranidalactivitiesareveryrarefbrthefirstday．Aftwexceptionshavebeenobservedinanestof V
cTYlb［V（C6601）intheearlynestingstage．Inthisnestthefbuowingbehaviourswereobservedbysevenworkers
onceitleaCllCaSe：（1）Re−CnlargementoftheceIl＼Va11＼＼∫ithoulIleSttllaterials（three、、・Ol−kers），（2）E一一Iarge−
me11tOrthecellwallwithnestmaterialscollectedbyherseばwithinthenest（OneWOrker），（3）Enlargemel宜
Oftheenvelopewithnestmaterialscollectedbyherselfwithinthenest（OneWOrker），（4）Malaxationofprey
withprovisioningtolarvae（OneWOrker），（5）Cleaningafbra由IgWOrkersoakedwithminintherield（One
WOrker）．
AsearlyastheirseconddaytheybegintoengageirlaVarietyofintranidalactivities，SuChasce11initia−
tion，CCllenlargement，PrO＼′isioningbroodandthermoregulationetc．，thoughtheyareabletobedislinguished
f和mOlderworkersbytheirbodycolour．Someofthemmaketheirfirstnightat2daysold，aSSOOnaS
theyareendowedwith flyingability．
3）Orientation flight and fbraging trips
ComparaモivebiologyorthenveJapanesespeciesorthegenus掩軍α（Hymenoptera，Ⅵspidae）
81
職ble28・Percentoftimespentineachactivityorasi喝1e掩軍α∫血f／加αWOrker（No．9）duringitslif如im已
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rromlarvae
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〓
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7
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鋸
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Ⅵ短ter collection
3
2
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一一一3・1
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一一一一一
一一一一一
Extranidalacl．
6
3
9
3 2．5
5
（llOur5）
Thefirstflightwasobservedat2to4daysoldin247individuauymarkedworkersofthenve挽野aSpeCies
（rrhble30）・Whenaworkerfirstleavesthenestshernakestwoorthreeorientationflightsduringthecourse
Ofwhichshelearnsthepositionofthenestentrancerelativetoneighbouringobjects．Theworkerfirst
niesstraightoutofthenestfbrabout20∼30cm，thenturnstofacetheentrance，flyingsIowlywithtoN
anふfromovernent．StiIlfacingtheentrancesheextendsgraduallythedistancefromthenesttosome3∼5
rnetres，andthenabruptlyreturnstothenest．Thesecondflightisusuallycarriedoutonthesameday
inasimilarmannertoもhefirst，butthedistancefromthenestisextendedto15∼20metres．This▼may
givetheworkersuffieientinformationthoughsornerequlreathirdorientationflight．Thefirstorienta−
tionmi帥ttook払rabout70−1005eC．andtheseconddecreasedto130−150sec．inmany掩軍αWOrkers．
ThefirstfbraglngtripISperfbrmedafterthesecondorthirdorientationflight−Duringthistriptheyoung
WOrkercouectsa11typesofloads，SuChasnestmaterials，Water，liquidfbodandprey，inasimi1arwayto
Older onesin the field．
4）Activitiesofmarkedworkersduringtheiradultlift
Theactivitiesofeachofthemarkedworkersofthefivespecies，duringtheiradultl脆aresummarized
in職bles28，29，Figs・54∼56・Theseresuitsindicatethattheworkersof挽甲atakeanactivepartinthe
affairsofthecoIonyverysoonafteremergence．Theybegantoperfbrmavarietyofintranidalactivities−
broodcare，neStCOnStruCtionandnestpreservation−aSearlyastheirsecondday・Theyforageda鮎ronly
82
Makoto MÅTSUURA
afewdays．Thereafter，bothextranidalandintranidalactivitieswerefbundinworkersofallageranges
andal10ftheworkersdailyperformedforagingactivitiesuntiltheirIastdayalive．Judgingfromthese
results，tOgetherwithmoreobservationsonotherindividuals，itiscertainthatadivisionoflabourbased
uponagedifferenceisnotestablishedin Va5PaWOrkers，unlikesome挽甲uぬworkersinwhichthereis
SOmeCOrrelationwiththeirage（EDW＾RDS1980fromPoTTER1964）・
AnotherimportantfぬtureintheproportionlngOftimespentinvariouskindsofactivityisthatI卸a
workersspentaboutonehalfoftheirtimeduringthedayrestingwithorwithoutslowwalkingonthe
comb．Theseunproductiveactivitiesareregardedasprovidingastand・byworkforcewhichcanberecruited
wheneverconditionschangeandthecolonyneedstomeetanewsituation．
SimilarresultshavebeenobtainedinApjsmeHif＞rvworkers，Whichspendabouttwo−thirdsoftheir
timeeitherrestingorwanderingthroughtheinteriorofthenest，anaCtivitythatLrNDÅUER（1952）refbrs
toaspatrouing．Heexplainedthatthesetwooutwardlyunproductiveactivitiesenhancedthecapacity
ofthecoIonyasawholetorespondtocaprlCiouschangesintheenvironment・Thisaccountcouldbe
appliedtothe掩5PaWOrkerstoo，andthedailylifeofworkersisingeneralarepetitionofthecyclecon−
sistingofresting，patrOilingandperformanceoftasksasstatcdforthehoneybee（SEKIGUCHrandSÅK＾G＾MI
1966）．
7hble29．Percentoftimespentineachactivityofasingle tespaanatisworker（No14）duringitslifbtime・
Ageindays
o12 3 4 5 6 7 8 9
Extranidalael．
− −
Orienla【ion
0．：
Pulpcollection − − −
Foodぐ01王ec【ion
−
Watercollection
−
Noloadcollec．＿
−
−
−
−
−
−
−
−
−
−
−
−
−
−
−
l・2 7・5 6・610・6 9・5 3t4 4・5 6‘8 5・2 6・0 3・5 7・2 5−5
−
−
3．1 9，3 5．9 5．816．Z11．2 5．314．3 7．3 11．0 4．9 7．8 5−8
− 王．2 1．11．ヱ 0．2 0．5 1．0
−
−
− 1．1 0．2
−
−
1．O
3．5 l．4 3．4 5．4 3．5 3．5 2．1 5．5 2．4 5．7 3．王 2．5 3．0
Intranidaiact．
Pulpmaはxation − −
1・2 2・4 3・2 4・5 8・3 7・8 2・5 6・0 7・5 6・Z 7・2 4・5 9・5 6・2 ロ㌫●
1．2
s。r。pi。g。，lVel。P。−
EnlarglngenVe． − −
Makingce11
の
コ
−
Making。edi。。1− −
托edinglarvae
4．13．12．】0．51．5 2．51．5 0．2 2．11．51．iO・3 3・4 2・5 蔓
2．2 6▲2 6．5 フ．212．111．】4．0 7．510・2 7・4 7−8 5t310・Z 8・0 冨
4・2 Z・4 3・8 2・11・5 Ⅰ・12・8 2・2 0・10・5l・る1・2 0・5 2t91・5 詔
0．10．2 0．l− 0．3 0・2 0・1− −
0・1− −
0・2 0・1冨
5．5 3．15．9 3・7 6・14・13・3 2・14・4 5・8 3t3 2・5 3・】610 5・2 旨
−
2．12．8 5．14．5 5．17．510．1】3．3 5．910・2 4・9 5・9 6・2 4・9
Preymaiaxalion −
紬
O・21・2 2・2 2・2 3・21・5 2・2 2・91・る 0・5 0・6 0・81・2 2・4 2・る 0−9
4・8 琵
11S
Foo
写霊誓三rYae 3・16・2 3・11・8 2・5 2・0 3t5 2・l】・5 0・5Ⅰ・1lt3 2・11・5 3・2 ＝
BodycIeanlng 5・2 】・10事5 0・3 0・2 】・0 0・5 0・6 0・3 l・0 0t5 0・3 0t5 0・3 0・3 0・6
1・1 0−3 1t3 2・2 0■5 0・6 0・4 0・4 1・1 0・2 0・4 0・】】・11・2 0・5
NesICleanlng −
Fannlng
− −
−
Spitli喝Water
0・2 0・3 0・5 0・6 0・5 0・3 0・2 0・2 0・1 0−1 0・3 0・3 0・5 0t5
0・2 0・3 0・3 0t2 0・2 0・3 0・2 0・3 0・4 0・1 0・3 0・5 0・4 0・6 0・4
Reslingwith
occasiona1 91．5 77．2 77．3 64．2 49．7 52．2 35．5 26．0 50．9 60．3 34．5 55．7 46．7 62．6 37．0 52．4
e亡11palrOl
DurObs●
3 3 3・5 4 2・5 8・5 9 4 3
Con叩araliveb圭0logyortlleriveJapanesespeciesorthegenus抱卵α（Hymenoptera，Ⅶspidae）
Fig．54．Activitiesoftwomarkedworkers（No．38：ロ，No．
41：田）of柁軍αぐ揮わ和duringtheirli托time（No．
38：17days；No．41：25days），
83
Fig，55．Activitiesoftwomarkedworkers（No．5：□；No．
6：畳）0＝収卿＝M／JdariTliaduringlhcirlirctinlC
（No．5：18days；No．6：22days）．
Fig．56．Activitiesof抽Omarkedworkers（No．2：□；No．3：慣）orIセ岬α什qpfcαduringtheirlifetime（No．
2：53days；No．3：61days）．
84
Makoto Mノ汀SUURÅ
一泡ble30．Agewhenrirstorientationnightsweremadeby物αWOrkers．
Daysarteremergence
2
3
3
4
3
5
8
︹B
︵U
0
3
Q′
11
7
l
′0
・l
・1
・l
00
仁JrqpJぐα
1
l
巨二／打打／Idロ′／／一由
5
00
一q
ト：ロJJロ／f∫
0bserved
5
7
Q′只︶
︼J
ト：ぐ和むJ・0
4
3
つー
卜：∫f〝JJ／／l〃J♂
No．or
individuals
一−▲
3．Division oflabourin an orphanedeolony
ItiswellknownthatwhenvesplneCOlonieslosetheirqueen，SOmeWOrkersbeginovlpOSitingwithinseveral
days（Marcha11896，lshay1964，Potter1965，Montagner1967），andthecolonycontinuesuntilthedisap−
pearanceofthelastworkers（MATSUUR＾1968）．However，nOdetailedaccountsofthedivisionoflabouramong
in orphanedcolonies has been pし1blished．
InthissectionsomefeaturesofthebehaviouroflaylngWOrkersaredescribedinrelationtothedivision
oflabouramongworkers．Thefbllowlnginformationismainlybasedonthedatafromanestof Vana始
（A67O5），inwhichthequeenwasremovedartificiallyonJuly24th，1967，andthen
daysafterremoval．Thequeenwasattackedandkilledbysomeworkersimmediatelyuponreturning・
1）Behaviour ofworkersand colon）Idevelopment
Disorganizationofthecolonybeganwithqueenremovalandsomeworkersfbughtoneanotheronthe
fouowingday．Fightingamongworkerscontinuedforthreedaysandthreeworkersleftandneveragaln
returnedtothenest．OnJuly29th，aWOrker（No．1）begantoovipositand，therea鮎r，thecolonyrecovered
thelostcohesiveness．Shebecamedominantinherrelationswiththeotherworkersand魚・equentlypersecuted
them（恥ble31）．Thisdominantworkerwastheonlyegg−layerandsurviveduntiladisorganizedstateof
thecolonyoccurredonSeptember28th・
Thble3l．DivisionoflabouramonglOmarkedworkersof tbpaana］isinanorphancolony（A6705）during
atotalof5Odays（70hoursobservation）afterthedeathofthequeen．Numbersgivethetotalnumber
Ortimeseachtaskwasper払rmed．
IndividualNo・
2
1
4
3
6
5
血eragein
7
Nos．ユ・10
9 10
8
28 45 29 3718 34 27 王8 29
t
sil
開
1
502
7・・・l
5 3 7
5 7
4 7
つ・ ▲フ︼ ′n
0U 4 00
3 5 1
5 Qノ 1
1 1 00
0
2456
つ︼︻／ 5
つ一つ︼ 1
8 つ一
0 8 4
﹁⊥
7 4 4
2
Food co】】亡Ction
Ⅵぬter collection
つ︼ 0ノ 1
Pulpco】lectioII
八U ■・1 8
0 ︵U ︵U
Extranidalact，
29・4
2
1
082
■・1
2 Q′0
7 0ノ
0 つ︼﹁・′0 0
．1 2 7 ■J 7
5
4
5 2
﹁−0 ﹁−
8 0ノ 2 1
4000505320190
4 へ∠
′O Q′ つJ 4 つ︼ 7 ′0 0
つJ 3 7
0
1
1 2 7 ハV ハリ
4 5
2 つ血・1
1 0
∧U
4 7 0
‘U
3 5 7 0 6 0 5
7 11
2
2 3 5
2
2
︵1．〇
1 5 3
0 0 0 0 ′人U ′0 9 0
つ〟
3 4
7
4
2 4 つ
J 3 5
8 莫U 2
7
7 3 0 2
′n︶
1 4 4
4 0 0 8
1
5 2 5 5 ′0 5 ハブ
Enlarsingcnvclope
Makir鳩CeII
Makingpedicel
Prey malaxation
托edinglarvae
Spittingwater
NesICleani11g
Oviposition
9
1ntra】1idalacl．
ComparativebiologyoFlheriveJapanesespeciesorthege】luS掩岬〟（Hymenoptera，Ⅶspidae）
85
Fig・57・ChangcincompositionorbroodandadultpopulationsinanorphancoIonyof tbpaana／is（A6705）
afterthedeathorthequeen，Arrowindicatesthedeathorthequeen．
Fig・57showsthefluctua亡ionsinthenumberofworkers，Cellsandofnestcontentsdurirlgtheperiod
fromremovlngthequeentothedisappearanceofthelastworkers．Themaximumnumberofworkers
reached580nAugust19th，andthereafterdecreased．Theimmaturestagesunderthequeenrightcondト
tionalireachedtheadultstageandauofthememergedasworkers．Therateofworkeremergencewas
2・lindividualsperdayonaverage，duringtheperiodn・OmJuiy26thtoSeptember4th，Whenthelast
WOrkeremerged．Ontheotherhand，themalesemergedfromSeptemberlstandthereaftercontinued
toemergeevenafterthedisappearanceofworkers．Itisassumedthatallthesemaleswereproducedfrom
eggsiaiddowllbyworkerNo．1．
Theratesofceuconstructionintheco主onywereconsiderablylowerthanthoseinqtleenrightcolonies
throughouttheobservationperiod（Fig．57）．Thetotalnumberofce11sconstructedafterdeathofthe
queenwasonly17，and801dcellsweredestroyedbyworkers．AtthelastobservationonSeptember30th，
the nest consisted oftwo combs with169cells．
2）Divisionorlabouramongworkers
Thble31givestheindividuaIdifferencesinfunctionamonglOmarkedworkerswhichremainedinthe
nestafterthedeathofthequeen．Asstatedintheprevioussection，WOrkerNo．1becameveryaggressive
andwasalwaysdominantinherirlteraCtionswiththeotherworkers．Shefrequentlysolicitedfromother
WOrkersaggressivelywithmouthparts．ThisworkerstoppedherextranidalactlVltyOnthedeathofthe
queenandspeciaiizedinovIPOSition，1．e．，Shebecamethesoleegg−1ayeramongtheworkersofthecolony．
Ai149examplesofovipositionobservedwereexcまusivelydonebyworkerNo．l．JudgingfromFig．57and
7hble31，WOrkerNo．10VIPOSitedO．73eggsperhourofobservationanditisestimatedthatatotalof
86
Makoto MÅTSUtjRA
4O8eggswerelaidin61days，anaVerageOf6．7eggsperday（maximum：24eggsonAugust24th）．Wbrker
No．1continuedaIsotomalaxatepreyforprovisionmgtothelarvaeaslongasshelived．
AllthemarkedworkersexceptfbrworkerNo．1，engagedinallextranida‡activitiesduringtheobserva−
tionperiod．ActivitiesindidethenestotherthanovIPOSition，SuChasnestconstruction，broodcareand
nestpreservation，WereSharedbyalIworkersinthecolonylnCludingworkerNo・l・
3）Characteristicsoflayingworkers
ThelaylngWOrkerNo．1inthecolonybecameeasilydistinguishablefromtheotherworkersonthebasis
ofherexternalfbatures，beingcharacterizedbythelossofbodyhairsrcsultinglnablackshiningbody．
TheseexternalftaturesofthelaylngWOrkerinanorphanedcolonyshowamarkedresemblancetothe
queeninpopulouscoloniesorthesamespeciesがterAugust（MÅTSUURÅ1968，1974），andthis“subs一山ト
tionqueenけwasfrequentlyobservedinorphannestsofthesamespeciestakenfromearlyAugusttoearly
No＼′ember（Thbles32and33）．
InthesesubstitutionqueensthelossofbodyhairswasconspICuOuSOnthegenaandmandibularbase，
andonthethoraxandabdomen，aSrepOrtedin V simi／〟mabyY＾MANE（1974）．
Therearesomedistinctdifferenceswithregardtotheovariandeve壬oprnentoftheworkersbetween
queenrigbtandqueenlesscolonies．Inthequeenrig机coloniesthemajOrityofworkershadundeveloped
ovarioleswithinclplentCOnStrictionsorwithdistinctconstrictionsmarkingtheappearanceofoocytes
（Fig，58），eXCeptOneOftheworkerswhichpossesseddevelopedovarieswithmatureoocytesincolony
A7521couectedattheendoftheseason．Ontheotherhand，inqueenlesscolonies，theproportionof
workerswithanumberofmatureoocytesasweuassuccessivelydevelopingones（Fig．58and職ble33）
wasverymuchhighercomparedtothatofworkersinqueenrightcolonies，andsomeofthempossessed
ovarieswithfu11ymaturedoocytesnotverydifftrentfromthatofqueens（Fig．58）．Inananalysisofovarioles，
thelargestqueenoocytewas3・4mmlongandthelargestworkeronewas3・3mmlong，andthenumber
ofovarioleswas7／7（Ie魚／ri帥t）inqueens（n＝12）and6／70r7／？inworkers（n＝35）・Furthermore，the
lbble3ユ．0、′ariandc、・・elopn−enL Or拍spa‘7na／iい、・Orkersin queen−rightcolonicsL
No．oFworkersineachstageor
e
e d
O C
S
N
110
62
ヘム
つー5 3
1 ∧U
つJ 3
4 4 4 0
くJ
0
1 3
O O O
■1 1
′0 ﹁J ︵U ︵b 8 0
O 八U 1 7
つー4
O O O O O Oノ
1144
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O
ハU
八じ八U ︿U O O −・1 0−1・1 2 0
O O
O
0
︵U
O
0
ハU
0
7
0
八じ
nU O O O O ︵U ︵U O
0
1
〇
八U
0
-P-
0
萱
〇
5
4
学
︵U
Ii III lV
0
Ⅰ
A6705 Jul．23I67 12
A7207 J山．30り2 10
A7510 A喝．i4’75 12
28
A7513 Aug．ヱ0’75 75
67
Å6709 A一喝．24’67 22 18
A6711 Sep．2’67 32 23
Å？404 Sep．2り4 18 37
A6712 Sep．4’67 35 65
Å7211 Sep．10り2 48 40
Å7212 Sep．13り2 32 27
Å7406 Sep．20り4 27
25
A7310 Sep．2㍗73 32 20
A731乏 Sep，23’73 43
40
A6820 Sep▲24’68 33 26
A73ユ3
Sep．24り3
7 13
A7108
Sep．25’71 63
45
Å7517 Sep．27り5 49 41
A6821 0（：t．る，68 48 77
Å7521 0（：t．7り5 10 26
A7215
0ct．10，72 10
28
一ヽ7110 0cl．18り1 9 ユ0
＊SeeFig．58
−恥talno．or
adul＝10rnetS
ovariande＼′e）opment．
Date
87
Compara【ivebiologyortileriveJapanesespeciesorthegenus物α（HymeI10ptera，ⅥさSpidae）
lbb）c3310＼∫ariande、，elopmcnt or［伽paarla／is、、■Orkersin queenLesscolonics．
Nest code Date
No．orworkersineachstageorovarian
development＊
l1
l
111
1＼／
4︵U
5
44
る3
1
3
爪U
4
′8
7
4
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1
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くJ ′0′O RV 7 83
ーハ凸
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2
．・′●＼
つム
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4
00
ハ
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︼ n70
4
1
33
＊SeeFjg．58
…ValueinparenthesesistllenumbeTOfsubstituteqtleerlS．
0
000n︶
1
︵U
3
38
2
3
4−1
00
ブーq′
つん
l
A7509 At唱．1り5
A6706 Ål鳩tlγ67
A7309 Åt鳩．20’73
A75ま1 Åug．ヱ0り5
A7512 Aug．20’75
A7308 Sep．12り3
Å？311 Sep．23ワ3
A7516 Sep．26り5
A7314 0ct．1，73
A7213 0ct．6り2
A7517 0く：t．7，75
A6904 0ct．10，るウ
A7216 0ct，12り2
A6717 0ct．14’67
A7111 0ct．19ワI
A7315 0ct．25，73
A7522 0ct．2ダ75
A6827 Nov．1’68
A7114 Nov．4，71
A6905 NoⅥ 8，69
7btalno．or
adul＝10rI】elS
︰巨．れ
1．■︰︰
頂鰯巌㈲臓紛潮目㌦〓
も亀
・
IV
Fig．58．Ovariesin挽御用皿血worker（Ⅰ一三Ⅴ）atldqueen（right）：トⅠⅠ；undeveloped，IlI；ratherdeveloped，
ⅠⅤ；developed，飴1lowinglos50fqueell．
numberoffullymaturedoocyteswas8to13inqueens（n＝13）and7to14inworkers（n＝35）．These
dataindicatethattherearefewdiffbrencesinthesizeandvolumeofovariesbetweenqueensandworkers，
IrlthequeenlesscoIonies，however，itisnotclearwhetheralltheworkerswithfullymaturedovarioles
COuldlayeggsorr10t，becausethesubstitutiorlqueeninthecolonyA6705wastheonlyindividuaまobserv−
edovipositing，andthenumberofsubstitutionqueenswasalwaysoneintheotherqueenlesscolonies
in the sa111eSPeCies（Tbble33）．
lnadditiontotheexternalfeatures，thesubstitutionqueenischaracterizedbyaremarkableprolonga−
tionofitslifespan，COmparedwiththatofworkersinthesamecolony．IncolonyA6705thesubstitution
queendiedonSeptember28th，thusshesurvivedfbrasmuchas67daysafterthedeathofthefounding
Makoto MATSUURA
88
職ble34．bngevitieso‖物四川胴血workersinanorphancolony（A6705）・
No．or
Lifespan（indays）
indi． Timeofdea【h
Obser．
≠brkerNo．1（Subs【ilutequCen）．
Workers，nOtlaying…
＞67
1
Sep・ヱ81h’67
露ニ2Z．3士S．d．18．5，Range：ユー45 41 FromJuly251h
10Sep．191ll
● DaleO（cmrgenceis uncertain．
＝Theseemerged fromlateJulytoearlyAugust．
queen・ThelongevityⅥ′aSabouttbreetimesaslongasthemeanlongevityortheworkerswl−ichemerged
fromlateJunetoearlyAugustinthesamecolony（rrhble34）．
4．Discussion
l）Behaviouralchangeorqueeninearlynestingstage
Oneofthemostcharacteristicfbaturesinthesocialinsectsisthefunctionaldivisionoflabourbetween
reproductivesandnon−rePrOductives．Thevespinewaspsaregenerallyconsideredasrepresentingthemost
advancedstageofsocialwasps，eXhibitingamarkeddifferentiationoftwofemalecastes，thequeen
largerandalmostexclusivelyegg−1ayingfbmale，andworkers−Smaller，Sterilefemalesandexclusively
haplometroticnestfbundation（atleastinso紘rstudiedtemperatespecies）・Throughoutthesoutaryperiod，
thequeenbehavesasifshewereasolitarywasp・Withtheemergenceofworkers，herbehaviourvaries
inbく）thquantityandqualitylnthecourseofcolonydevelopment・
Inthevespinewasps，VERLA靂NE（1932）hasobservedin伽′∫cカov朗p〟ね椚ed由thatthequeenwouldcon−
tinuetofbrageandbuildanesttwomonthsartertheemergenceortherirstworkeちbutinthisco10ny
thenumberofadultswasonlytwelve．BRIANandBRIAN（1952）reportedslackeningofqueenforagingof
Dolichove5Puぬ砂／vestrissoonaftertheworkersstartedfbragmg，althoughitisqtlitepossiblethatshe
hadreducedheroutdooractivityearlier；Shewasnotseentogooutafterthe33rdday，butonequeen，
whichproducedonlythreeworkersfrequentlyleftthenestaslateasthe50thdayandthischangeispro−
bablyrelatedinsomewaytothepresenceoffbraglngWOrkers．In掩軍uhtgermanjcaand Vv塘ariswith
populouscolonies，SpRADBERY（1965）statedthecessationofextraovipositionaldutiesbythequeenwhen
fburormoreworkerswerepresent．SHiD＾（1959）observedin陀sputaj7avicqpsthatthequeenhadceased
toleavethenestwith11workcrsonJunellth．AstoI包甲aSPeCies，IsH＾Y（1964）reportedon Vorienta胎
thattheworkersaidedthequeeninnesteonstructionandlarvalnursingbetweenmid−MayandmidJune
inIsrael，andthequeen，then，PrlnClpallyoccupiedherselfinovlpOSitionwithoutgolngOut・In Vana〟ち
aqueenstoppedherextranidalactivitieswhensixworkerswerepresentinearlyJulythoughshestilicon−
tinuedsomeintranidalactivities（M＾TSUURA1974）．
Thepreviousreportshavedealtalmostexclusivelywithforagingbehaviourofthequeen・Littleatten−
tionhassofarbeenpaidtointranidalactivitiesofthequeenintheearlynestlngStageS・Evenafterthe
queenof物aceasestoleaveanest，Shecontinuestoengageininsidework，i・e・Ce11enlargement，PrOVi−
sionofrnalaxatedpreyandliquidtothelarvaeandthermoregulation，fbraboutamonth・Thus，the
cooperativenestingstagebetweenqueenandworkerslastsfbrabout5Odaysaftertheemergenceofthe
firstworkerinthenestof Vc7Vb7nOntheotherhand，in掩軍alrqpIcqwhichhasthesmallestcolony
sizeamongtheIセ軍αSpeCiesinJapan，genera11ysimilartothatortheJapanese伽触e∫incolonysize
（MATSUURA1970），thequeenparticipatesinintranidalactivitiesnotonlysoonafteremergenceofworkers
butthroughoutherlife（MATSUURA1974）．Thesefactssuggestapossiblecorrelationbetweencolonysize
andgradualchangeofqueenactivitie＄in物aspeciesinsuchawaythatthelargerthetypicalcoIony
sizeofaspecies，theearlierthisoccurs，thoughsuchchangeisobviouslyrelatedinsomewaytothepresence
Of≠・Orkers，CSPCCia11y Foragers．
Comparativebio10gyOfthcfiveJapanesespeciesofthegenus掩即a（Hymenoptera，Vbsp主dae）
89
工npolistinewaspsthetasksofcoIony−buildingandbroodcarearesharedbythefoundressandworkers，
thoughthefbrmerisusua11ythesoleegg−1ayerandshehasatleastfburdistinctivebehaviouralcharacteristics：
egg−laying，Superiordominance，Cellinitiation，andspecializationinpulpforaging（Ⅵ短st−Everhard1969）．
West−Eberhard pointed out that cellinitiation and pulp fbraging by the fbundress of Polistesare
characteristicsdirectlyassociatedwithdevelopedovariesordominanceasshownbytheexperiments
PARDl（1948）．In紬触eshuntterいhefbundress．takesamassofpulpfromareturningworkertocon−
StruCtneWCellsattheedgeofthecomb（MzcHENERandMICHENER1951）．
Ontheotherhand，nO掩申αqueenSeng喝eincellinitiationarterworkereme柑enCe，eVenin H胱画路
Theabandonmentofcellinitiationbythequeenisprobablycorrelatedwith−1argercoIonysize／Evenin
thecooperativeperiod，themodeoflifbin陀甲aiSregardedasm6redifferentiatedthaninpolistinewasps．
Furthermorethisperiodmayrenectaprlmitivepolyethismbetweenqueenandworkersintheearlierstage
Ofsociale＼・Olutionpassedthroughby柁spqspecies．
2）Relations betweenlabourand agein＼＼・Orkers
Age−COnditionedpoiyetbismhasbeenrecognizedamo喝theworkersinacolonyofthehoneybee（R8scH
1925）andants（SuDD1967）・Insocialwasps，pOlyethismaccordingto喝eisweaklydefined．紬／如e∫ノα拡
WLgaeWOrkersdonotfbrageuntilaboutaweekfbllowlngemergenCeandtheolderworkershaveagreater
tendencytoparticipateindomesticwork，butthereisotherwisenoclearschedulingoftasks（YosHIX＾WÅ
1963）・In掩spuぬspecies，Shida（1959）reportedin VPavicqpsthatcollectingnestmaterialswaschiefly
Performedbyolderworkers．AccordingtoMoNTÅGNER（1967），挽甲utaWOrkersassistinnestbuildingduring
【herirst托wdaysartereme柑enCe，andthentbeyalsobe蛮ntofbedlarvae；rinally，SOmeindividuakadd
foragingtotheirrepertory．BRlÅNandBRZ洒（1952）statedthattheworkersofDo〟chove5Puね砂Ivestrね
assistedinthemalaxationo‖1iesbrou8加inbyotherworkersverysoonarteremergenceand払raged
Onlyafewdaysafteremergence．GAUL（1948）fbundinhisobservationsonsomecoIoniesof掩3Puねand
Dolichovepulathatnoagedis
1nthecaseofle5Pa，aSimilarpatternhasbeenrecognizedandthe、、′Orkerstakeanactivepartin、′arious
tasksoFthecolon〉・Ver）・SOOn afteremergence．
Ontheotherhand，intheworkersorthehoneybee，バタね沼e／／推喝SOmeaCtivitiesdependonparticular
physiologlCalconditionswhichareage−COnditioned；forinstance，fboddischargefromthehypopharyngeal
glandsandwaxsecretiononwaxglands，etC．（KING1933）．In tespaspeciesitislikelythattheworkers
havenoschedulesfornurslngandnestconstructionwhicharelimitedbyglandualractivityasisknown
inthehoneybee．Therefbre，itappearsthat掩spaworkerscanperfbrmanytasksaftwdaysafteremergence．
3）DivisionoflabourinanorphanedcoIony
Inthecourseofcolonydevelopmentaconsiderablenumberofcolonieslosetheirqueensbyaccident
Ornaturaldeathaftertheemergenceofworkers．Nevertheless，mOStOftheseorphancoloniessurvivefor
rnorethanafbwmonthsand．frequentlycontributetomaleproduction．
MÅRCl寸AL（1896）firstreportedinhisstudiesonIセspu血spp．thatworkersdevelopedtheirovariessoon
afterremovingthequeen，andsomeofthembeganovlpOSitingwithinlOdays．In掩spaalso，itisobserv−
edthatworkersovipositedseveraldaysafterthequeen，sdeath（IsHAYetal．1965，MÅTSUURÅ1968）．These
CaSeSindicatethatsomeworkersofvesplneSpeCieshavethepotentialabilitytoovlpOSitintheorphaned
COnditioninspiteoftheirdifftrencesinmorphologicalandphysiological魚ctors（TAXAMATSU1952，BLÅCKrrH
1958，SpR＾DBERY1965）．Inthecaseof VanalLfitisrecognizedthatsuchlayingworkersaredifferent
n．omtheotherworkersinorphanedcoIoniesandarecharacterizedbythefollowingpoints：（1）theprimary
OrSOleegg−la〉′er，（2）10SSOfhairsonthegenaandmandibularbaseandpolishedhead，（3）cessationof
allextranidalactivities，（4）themostsuperiordominance，（5）receivingagreatdealoffbod．fromtheother
WOrkers，（6）remarkableprolongationoflifbspan．Ofthesecharacteristic，（1）（2）and（3）arecommon
tothoseofthequeenof柁軍aSPeCiesinapopulouscolonyiandagonisticbehaviourinvoIvingdominanee
relationshipsisvirtua11yabsentbetweenqueenandworkers，nOtOnlyatfbodexchangebutthroughout
theirencounters．Ontheotherhand，incoloniesofthemoresociallyprimitivevespidwasps，SuChas
MLgchoqttarusdnweniand撤触esspp．，thedivisionoflabourbetweenqueenandworkersISPrlnClpally
90
Makoto MATSUURA
broughtaboutbybehaviouraldominance（YosHlXAWÅ1963，MoRIMOTO1961，WEST−EBERHARD1969，JEANNE
1972）．Asirnilarcaseisobservedamongfbundressesofjb撤tesspp．intheprimarypleometroticcolonies
OfPa［lmL／arLs（R＾U1940），Pga［／ictJ∫（P∧RDl1943，DELEUR＾NぐLi1955）andPveJ苫icoIor（RtcH＾RDS1971）．
InthesespeciesthecoloniesarefbundedbyanumberofalmostequlPOtential，mOStlyfbrtilizedfemales，
andtheonewiththemostdevelopedovariesbecomesthequeenwhichforagesmuchlessthanthesuboト
dinatesorworkersandreceivesagooddealoffbodfromthem；theotherseitherbecomeactingworkers
Withovarianregressionorleavethenestasaresultofcompetition．
1n物aallworkersinqueenィightcoIoniesseemtobealmostentirelyequipotentialinreproductive
functiorlSCOmparedwiththatofthequeen．Theconditionsintheabsenceofaqueeninthecolonyof
H〝〃α梅beingofasmallco10nySize，areVerySimilarto抽oseinapleometroticcolonyoぎsome伽／如郎
spp．asabovementioned．Therefbre，itseemslikelythatthedominantworkerofVanalLscontroIssubor−
dinatesandpreventsthedevelopmentofovariesinsubordinates，Whichinvolvesalltheworkerswhoemerged
afterthedeathofthequeen．
However，Verylittleaggressiveoragonisticalbehaviourwasseenaftertheestablishmentofthesubstitu−
tionqueen，Whichbecamecharacterizedbythepolishedheadandthelossofbodyhairslikethequeen
Orthespecies．
V．POPULITlON DYNAMICS
Hornetsofthegenus物αpOSSeSSaCOmplexmodeorli托，andtheirannualcycleisfundamentally
dividedintotwobasicphases，SOlitaryandsocial，eaChofwhichoccupleSabouthalfayear・Therefore，
varioustheoriesofpopulationecologywhichhavebeenappliedtomostconventionalanimalpopulations
arenotapplicabletothecaseof掩3PaSpeCieswithoutappropriatemodifications・
Inthischapter，thepopulationof物aspeciesistreatedattwolevels，individualandcolony・Atthe
individua11evelthesurvivorshipofcolonymembersismainlydiscussedinrelationtothelifttabiesobtained
frornworkersinnormallydevelopedcoIonies・AtthecolonylevelIdealwithsomeobservationsonthesur−
vivalofnests，COlonydevelopmentandannualfiuctuationsofnestsandsomefactorsaffbctingthem・
1．Survivorshipcurvesof噸αWOrkers
l）Length oFde＼′e10Pmentalstage
Prhble35showsthevariabilityofeachstageandthetotaldurationofbrooddevelopmentinworkersof
籠ble35．kn紳OreaChdevelopmentalstageinfive挽甲αWOrkers（inday5）andthesizeoradultworkers・
Larval instar
Species Egg
H∫わ肋
Hぐ相加
H〝〃鵡
H〝榔血加血ぬ
H岬加
1Il 王ⅠⅠIV V
（）（（，（）（，（，
（，（，（）（）（）（，
（，，（，（）（（）
（（）（），（（，
（（）（）（，（）（）
Parenthesiヱednumbersshowstandarddeviations（土）．
● h／1ean or10＼＼・Orkers forcaeh speeies．
No．or
Size of
adult
observed workers＊（mm）
トV Cocoon 7btalindivid．
‖5
21・3
73
24・1
60
24・5
84
34・2
71
30・1
ComparativebiologyofthefiveJapanesespeciesoftheさenuS沌spa（Hymenoptera，Ⅶspidae）
91
fiveIセ3PaSpeCies・Comparlngthedata，thelengthsofthe5thinstarlarvalstagevarywidely，Whilethe
lengthsofegg，immaturelarvalstage，eSPeCiallylst−3rdinstars，andcocoonstagesarefairlyconstarltin
eachspecies，thoughthereseemstobesomespecificdifferences・7btaldurationofbrooddevelopment
fromeggtoadt11tvariedontheaveragefrom30．8daysinIてsimil［imainthesmauestspeciesinbody
Sizeto40・1daysin Vmandarifliainthelargestspecies，anditisrccognizedthatthereisatendencythat
thelargerthebodysizeinadulthornets，thelongerthedurationbecomesamongthefivespecies．
2）SurvivorshipcurvesforalldeveIopmentalstages
Figs■59∼63presentsurvivorshipcurvesねralldeveiopmentalstagesorworkersorrive挽甲αSpeCies，
usingthefollowingparameters：（1）lx，numberofindividualsaliveatthebeginningofeachstage，（2）dx，
numberofindividualsfailingduringeachstage，（3）qx，nurnberofindividualsfailirlgPerlOOOaliveat
thebeginnlngOfeachstage．Althoughallthesedatawerepreparedfromweudevelopedcoloniesatpolyethic
Period，thesefivespeciesshowagenera王correlationbetweensurvivorshipandmortality，andallthesuト
Vivorshipcurvesindicatebasica11ysimilarftatures，i．e”remarkablylowmortalityduringtheimmature
︵00−×︶
SJくっロー≧云竺︻
隻u
二望
∧G E IN D∧Y S
Fig．59．SurvivorshipcurvejbrI句卯Slmi／limaworkersinanest（S6801）．rrA：teneraladtllt．Forlx，dxand
qX，See teXt．
︵〇〇lエ
S﹂く⊃○︻＞︻⊇丁〓］○
10
20
30
40
50
dO
70
A G E 王 N D A Y S
Fig．60．Survivorshipcurveror掩叩〟紺地澗WOrker5inanest（C6601）．m：teneraladult．‡もrlx，dxand
qX，See teXt．
Makoto MATSUURA
92
▲8
′O
lOC■エs﹂＜コq ；完︻ ≡
0
■月
山〇．〇Z
10
20
30
dO
50
る0
70
A G E IN D A Y S
Fig．61．Survivorshipcurvefbr物aanatiSworkersinanest（A7207）．TÅ：teneraladult．Fbrlx，dxandqx，SCeteXt・
︵〇〇一ズ︶S﹂くっ凸T＞−凸こ︼
﹂〇．〇〓
A G E IN D A Y S
Fig．62．SurvivorshipcurveforI々spamar7dari17jaworkersinancst（M7401）／rA：teneraladult．Forlx，dxand
qX，See teXt．
stagesandhighmortalityafterthecommeriCementOfforaglngaCtivitiesintheadultstage．Therefore，
thesurvivorshipcurveisaconvextype，andmaybeconsideredtobecurve7ypeIinDEEVEY’ssense（DEEVEY
1947）．
3）Survivorshipcurvesぬradultwol◆kers
Fig．64showssurvivorshipandmortalitycurvesforthesamecohortsglVeninFigs．59∼63，butdeals
withadultsaloneatdailyintervals．Survivorshipcurvesforeachspeciesarecloselysimilarandingeneral
themaximumlongevityisabouttwiceasIongasthemeanlongevity，thoughitisrecognizedthatthere
aresomedifferencesamongthem．Themortalityof V trqpLCaWOrkersduringtheearlyadultstageis
remarkablyまowascomparedwiththatoftheotherfburspecies．Thisdifferenceseemstobeduetothe
diffbrcncesintheirfeedinghabitsanddispositionasdiscussedlater．Agradualdecreasebeginswiththe
ComparativebiologyofthefiveJapanesespeciesofthegenus掩spa（Hymenoptera，Vtspidae）
93
8
′0
ち○■エs﹂くっ凸︻＞︼雪〓
0
﹂〇．〇Z
20
10
30
40
50
る0
70
80
90
A G E IN D A Y S
Fig・63・Survivorshipcurvefbr ti5PatrqPicaworkersinanest（T7401）．TA：tenCraladult．Fbrlx，dxandqx，SeeteXt．
8
でUO︻×︸﹂く⊃q；−望三
0
S
h岩．〇之
10
20
30
50
dO
A G E iN D A Y S
Fig．64．Survivorshipcurvesforadultworkersoffive掩spaspeciesatdailyintervals．sim：VsimiIlima；man；
巨：／〃〝／Jr血／血血こCJV：レ：（・rロわrr）；〝冊：ト：〟Jlβ／J∫；rrO：lノ：什叩Jm．
COmmenCementOfthefirstextranidaiactivity2or3daysaftertheiremergenceinallcurvesexceptfbr
Vlr叩LC玖andsurvivalcurvesdescendwithasigmoidshapein eachspecies．
4）fもteorbroodatemergenceofsexuals
Themalesandnewqueensareproducedtowardstheendoftheseasonalcycle．Theyfbedonthesecretion
Ofthe5thinstarlarvaeandthefoodbroughtinbytheworkersduringthetimetheyrernaininthenest．
However，ithasbecnobservedthatmanydeadorweakenedlarvaearetakenoutoftheceusandabandorl−
edonthegroundneartlleneSt，With theappearanceofthesexuals（DuNCÅN1939）．
1bble36sho≠●S thelifctables fbrall111embers orthcbroodoral／：（l−OPiぐaCOlony（T7401），＼＼′hichis
Classifiedintothreeperiodaccordingtothetimewhentheeggswerelaid；SOlitaryperiod，CO−Operative
Periodsbefbreemergenceofsexualsandafteremergenceofsexuals．Thesedataindicatethatbrooddevelop−
mentis clearly disturbed towards thelater stages ofthe seasonalcycle．In the case ofthis nest，the
94
Makoto MATSUU】ミA
lbble36．Lifttable rorlセ5Pa（rOPicoin onenesl（T7401）at diffbrent nesting periods．
＊＊
＊＊
0
0
Qノ
2
〇一 0・4
7ノ
八nU
〇一
32 3 2 3 2
1
1
1
‖
′0
2
000
2
m
′h︶
Ⅳ
ÅV
V
（n＝269）＊
0
つJ
1
Eggs iaid
July14廿l−
befbre
emergenceor Aug．23rd
SeXuals
Cannibalism
by adults
Cannibalism
byadults
0
0ノ
Cocoon
Adult
0
（
0
Qノ
（n；＝39）＊
0
つJ
Qノ
lV
instar
0
d7
3
Lar＼・al
0
ハブ
m
June9th ∼
July13tIl
0
3
‖
Eggs laid
befbre
emergenceor
WOrkers
●＊
0
0
Qノ
3
Egg
q
和x＊＊
X
リー’しし‘1しq▲
X
Ju
Duration of
ovOn
X
Period
′0
Can】1iba】isI11
6
d
n
O h
O u
C
by adults
Cannibalism
by adults
CO
A
Ca11nibalis111
by adults
Egg
王
Eggs laid
a地r
Aug．24th−
emergenceor Sep．25tb
sexuals
I王
98
28
Starvation
Starvation
70
2
28．6
2．8
68
3
4．3
Star＼・alioll
65
6
9．2
Starvation
59
7
】l．S
52
47
5
5
brヽ・al
instar
111
IV
（Il＝98）
Cocoon
Starvation and
unknown
Cannibalism by
adults
Adult
90．2
100．0
0
＊NoofeggsovIPOSitedbythequeenduringeachperiod．
＊＊ See text．
mortalityofbroodwaszeroduringthesolitaryperiod．Highmortalityoccuredduringthetimeofthe
5thinstarlarvaeamdcocoonsartermid−August．
1tisclear什omcontinuousobservationsthatthemortalityfactorsinthedevelopmentalstagesduring
thelaterperiodwerealmostentirelyduetocannibalismbyworkersandthcsacrificedlarvaeandpupae
WeremOStlythoseofmalesornewqueens，Thiscannibalismisassumedtobecausedbytheiackofpro−
teinfoodwhichalmostentirelydependsonpolistinebrood，becausethecoIoniesofpolistinewaspshave
alrcad）・CeaSed theirbrood rearingarter mid−August（Fig．41）．
Duringtheperiodafteremergenceofsexualshighermortalitywasobservedthroughouta11devcIopmental
stages，particuIarly，eggand5thinstarlarvalstages，andnoeggswerelaidaftertheemergenceofthe
ComparativebiologyofthefiveJapanesespeciesofthegenus7bpa（Hymenoptera，鳩spidae）
95
SeXuals・Inthisperiodthemortalityintheearlydeve王opmentalstagesfromeggto3rdinstarlarva、and
COCOOnStageWaSmainlyduetocannibalismbyworkers・Butitshouldbenotedthatthehighmortality
Ofthe5thinstarlarvaewasmostlycausedbystarvation，nOtbycarmibalismaswasobservedatthetime
beforeemergenceofthesexuals．
職ble37isalifbtableforbroodofVana胎atthelaterstagesoftheseasonalcycle，anda11thebrood
inthecoIony（A7207）wereidentifiedaseitherrnadeornewqueen．lnthisspeciesatendencysimilarto
職t）le37・Lifetablefor掩5Paanatjsbroodfromeggslaidbythequeenbefbreandafteremergenceofthe
SeXualsin one】1eSt（T7ヱ07）．
Devental
period
肘＊
（
V
（36♀♀27杏容）
Sand
Cannibalism
42
3
39 92．8
43・9揖2・202・3
1
Cocoon
Adult
0
v
（n＝82）
1
Cannibalism
2
III
IV
3
Larval
inslar
つJ
Sep．25th∼
emergence of Nov. lOth
SeXuals
a魚er
14一
Eggs iaid
3
ll
CannibalisTn
Cannibalism
Cannibalism
I
6
Egg
35
Cocoon
Adult
2
（n＝124）
Cannibalism
Cannibalism
Cannibalism
Cannibalism
6
l＼／
・9
7 2胴一
・・0
51
703
1L・
111
insはr
1
Larval
0
Sep・24th
2
eme柑enCeOr Aug・12th−
7
befbre
5eXua15
Cannibalism
＝
117115115114112
2874‘
74U
6
34
4 4
I
Eggs】aid
闇
Cannibalismby
adults
Egg
3 100．O
O
＊ See text．
VtrppLCaisalsorecognized・AbouthalfofI24eggslaidftomAugust12thtoSeptember24thbeft）re
emergenceofthesexualsreachedtheadultstageand36newqueensand27malesemergedanddeparted
fromthenest∴Highmortalityoccurredparticularlyinthe5thlarvalandcocoonstages，andmostofthem
died丘omcannibalismbyworkers・Ontheotherhand，43・9甲boftheeggslaidbythequeenfromSeptember
25thtoNovemberlOthafteremergenceofthesexualdisappearedduringtheeggstageduetoeannibalism
byadulthornets，but47・6％ofthemdiedfromstarvationtowardstheendofthecoIonycycle．Onlythree
reacbedtbecocoonstagebu【noadultseme喝edinthまsperiod．
Judgingfrorntheseobservations，tOgetherwithotherobservationsinmanyothernests，itisconsidered
thattheworkersdevotetheirattentionmainlytonursingofadultsexualswhichhaveernergedearlier；
thusmostofthelarvaefailtobefedbecauseofadecreaseinattentionglVentOthemwhenthesexuals
emerge・Thisbehaviourofworkersmaybecausedbytheextremeclaimfbrfbodsupplybytheadultsexuals．
2・Seasonalchangesinthepopulationstructureo＝物卯COlonies
l）Number ofce11s
Fig・65showstheseasonalchamgesinthenumberorcellsinthecoloniesof汽veIセ軍ぽSpeCiesexamined
during1966∼1975．
96
Makoto M入TSUURA
f（trqpM：ThehibernatedqueensstartedtobuildtheirnestsinearlyJune，andmostofthecolonies
10SttheirqueeninSeptemberandnestactivitiesceasedinaboutearlyOctober．一幅oqueennestcollected
inmid−Junehad16and23cells．Themeannumberofceuspercolonywas37．6土S．d．13．0（n＝5）in
June，increasing to128．O j＝S．d．62．1（n＝12）in August and165．8土S．d．77．3（n＝10）in Scpternber，
respectively．
ハ00rるトS山Z∝山むSJJ山U
JO
■芝
Fig．65．Seasonalchangesillthenun】bcrorcellsintheeoloniesorfi＼・・Clセ5Pa5PeCic5．OpeneircIc．queenle5S
COlony；SOlidcircle，queen−rightcoIony；largeopencircle，rnOnth王ymeanwithstandarddeviation．
y：a朋Hs：ThequeensbegantheirbuildingactivitylnearlytolateMayandthecolonieslastedabout
Sixmonthsunti11ateOctobertoearlyNovember，though65．4勒ofsurvIVlngneStShadiosttheirquee
among26nestssarnpledinOctober．ThemeannumberofceIIspercolonychangedn・Om69．1土S．d．
54．8（n＝15）inJulyto516．7d＝S．d．190．1（n＝7）inearlyNovember．Queenlesscoloniesincreasednoticeabiy
afterearlyAugustand47．8％ofallthenestswhichwerecouectedfromAugusttoNovemberhadlost
theirqueems．
Vc用bro：ThequecnsbegantheirnestingactlVltylnmid−May．rrb［OqueenneStSCOliectedinlateMay
had29and31cells，reSPeCtively．ThenumberofcellsremainedlessthanaboutlOOuntilmid−Juiy，and
thenincreased．Themeannumberofcellspercolonychangedftom767．6土S．d．962．5（n＝7）inAugust
101，984．8土S．d．1，116．2（n＝＝6）illOctober．
f（mandarinia：QueensprobablybegantoestablishtheirnestsinearlyMaytoJune，andthenesting
activityceasedinlateNovember．Thequeennestscontained20celisinearlyMayand40ceusinlate
June．ThenumberofcellsincrcasedwiththeemergenceofworkersafterJuly；andthemeannumberof
ComparativebioIogyofthefiveJapanesespeciesofthegenus挽甲a（Hymenoptera，鳩spidae）
97
Cellspercolonychangedfrom284．0土S．d．239．O（n＝2）inJulyto2，712．4d＝S．d．984．7（n＝15）inOctober．
Vskn肋a：Queennestscorltained14．7ceus土S．d．2．1（n＝3）inlateMay．Thenumberofce11sin−
CreaSedgreatlyduringearlyJuly，andincreasedagainafterJulywiththeachievementofrelocationof
thenestsite．NestingactivitieswereobservedeveninearlyDecemberinsomenests．Themeannumber
Ofcellspercolonychangedfrom575．7j＝S．d．435．6（n＝22）inJulyto6，087士S．d．2，869．3（n＝14）in
October．
Themaximumnumberofce11spercolonyso如observedvariedfrom313in析′r叩JCαtOlO，058in
Vsimi〟ima（7hble38），andthenumberofcellsproducedisrankedassimil〟ma＞mandaT・inia≧crabro
夢α〝α胎＞ ∫rqpJC乱
Thble381Thenumberorcellsinthelargestnestso［rivelセ5PaSPeCicssor乙rtakeninandnear7（ibi．
Species
Nov．23，64
l
y．正閏沼血相
0ct．ユ2り3
Nov．21，64
313
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4，563
4，661
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Sep．18’71
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2）Im汀iatureStageS
Themeannumberofirnmaturesin物acoloniesaresummarizedinFigs．66∼70．
1て（ropica：ThenumberofeggsremainedatlO−20duringJuneandJuly，reaChedamaximumof28．2
土S・d・14・9（n＝7）inAugust，andeggproductioninmostcoloniesendedbymid−September．
ThenumberoflarvaewasalsohighestduringAugustandthemeannumberoflarvaewas25，7土S．d．
12．5（n＝7）forlst−4thinstarsand2l．3j＝S．d．18．1（n＝7）fbr5thinstarinthismonth．Fewerlarvaeof
lst∼4thinstarswerefoundinmostcoloniesinSeptember，butthe5thinstarlarvaeremainedrelatively
eons【ant during Septernber．
Workercocoon＄increasedfromOinJuneto15．6土S．d．14．2（n＝7）inAugust，anddisappearedin
Septemberinmostcolonies．Therneannumberofcocoonsofsexualswas18．7土S．d．23．7（n＝10）fbr
queensarld6．7d＝S．d，11．5（n＝7）formalesinAugust，andwas7．4土S．d．16．7（n＝11）forqueensand
4．5d＝S．d．7．3（n＝10）formalesinSeptember．
Vc用bro：ThemeannumberofeggspercoIonyremainedlessthan32untilJuly，andthenreached
amaximumof152．Oj＝S．d．138．2（n＝13）inSeptember．RapiddeclineoccuredinearlyOctoberandfew
eggs＼＼′ere foundin most coloniesinmid・October．
ThenumberoflarvaewasalsohighestinSeptemberandthemeanwas174．6j＝S．d．167．1（n＝13）for
lst∼4thinstarsand239．1土S．d．209．7（n＝13）for5thinstarinSeptember．Numbersoflarvaeofallin−
StarSdecreasedrapidiyaftermid−October．ThemaximumnumberoflarvaepercolonyinSeptemberwas
668fbrlst∼4thinstarsand719for5thinstarinanest（C7504）collectedonSeptember9th，1975．
Coeoons、、′erefbundinlownumbers（2−27）untilJuly．Themeannumberof＼＼・Orkercocoons＼＼∫aS123．7
土S．d．106．3（n＝7）inAugustwithamaximumof335inanest（C7304），anddeclinedduringSeptember．
ColoniescontainedrlOWOrkercocoonsinOctober．CocoonsofsexualsfirstappearedinlateAugustin
afbwcoIonies，andthepeakpopulations，Whichwerefb11ndwithmeannumberof85．2d＝S．d．133．2（n＝10）
fornewqueenand185．9土S．d．275．5formaleinSeptember，declinedrapidlytowardOctober．Themaxr
imurnnurnberofcocoonsofsexualspercolonywas420fornewqueen（C7507）and885fbrmale（C7504）．
l′：mL7（Jdar［nia：Themeannumberofeggspercolonyincreased from38（h／17401）inJul〉・tO176．8土
S．d．120．7（n＝16）inSeptemberandthereafterdeclinedduringOctoberanddisappearedinlateNovember．
DuringAugustnumbersoflarvaewereprobablyhighestineachstage．Immaturesoflst∼4thinstarsdecreas−
edthereafteranddisappearedbylateNovembenThemeannumberof5thinstarlarvaeinSeptember
remained234．3d＝S．d．162．4（n＝6）inAugustanddecrcasedgradualtyduringNovember．Themaximum
Makoto MATSUURA
50
dO
30
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Fig．66．Seasonalchangeinmonthlymeannumberofimmaturestagesincoloniesof物atr・qPica．Ⅶrtica1
AO
1ineshows standard deviation．
粥
20
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−○−1sT」廿日lトISTARS
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Fig，67．Seasonalchangeinmonthlymeanmlmberofimmaturestagesincoloniesof噸ac用bro，Ⅵrtical
主ineshows standard deviation．
ComparativebiologyortheriveJapanesespecie50rtheさenuS挽研α（＝ymenoptera，Ⅵ華pidae）
99
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Fig・68・Seasonalchangeinmonthlymeannumberofimmaturestagesinco10niesofIセspamanda7・inia．Ⅶr−
ticalline shows st8ndard deviation．
MAY Ju N JU L AUG S∈P O CT NOV
Fig．69．Seasonalchangeinmonthlymeannumberofimmaturestagesinco主oniesof物aana！is．Ⅵきrtical
lineshows standard deviatiorl．
Makoto MA・rSUU‡iA
叩
08
00
00
000
00
芸3旨㌧0広
00
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00
00
00
00
8
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Fig．70．Seasonalchangeinmonthlymeannlユmt）erOfimmaturestagesincoloniesof Vb3PaSimi［lima．Vertical
lirle Shows standard deviation．
numberoflarvaepercolonywas452（M7201）fbrlst∼4thinstarsand637（M7202）fbr5thinstarinSeptember
inanest（M7201）collectedonSeptember19th，1972．
Thenumberof＼、′Orkercocoonschanged from95．0土S．d．96．2inJulyto337．2士S．d．286．7（n＝16）
inSeptemberandthereafterdeclinedrapidly，disappearlnginNovemberinmostcolonies．Cocoonso
SeXuaJsrirstappearedinlateAugustina fewcoIonies．Thenumberormalecocoonsreachedapeakin
Octoberandthatofnewqueenwas88．3土S．d．130．1（n＝16）inNovember；bothdisappearedinDecember．
Themaximumnumberofworker，maleandnewqueencocoonspercolonywasl，O15（M7201）inmid−
September，532（M6902）inmid−Octoberand491（M7503）inearlyNovember，reSPeCtively．
f：anaHs：Themeannumberofeggspercolonydidnotexceedca．10untilearlyJulyandthereafter
increasedfrom18．O土S．d．15．0（n＝6）inlateJulyto42．2j＝S．d．36．8（n＝9）inearlyOctober，With
maximumof99inanest（A67O6）collectedonAugust17th，1967．Thenumberofeggsdeclinedrapidly
duringmid−Octoberand early November．
Thenumberofimmaturelarvalstagesoflst−4thinstarschangedapproximatelyequallytothatof
eggsthroughoutthenestingperiod，andthemaximumnumberofirnrnaturelarvalstagespercolonywas
89inanest（A6821）co11ectedinearlyOctober．Themeannurnberof5thinstarlarvaeremainedIessthan
thatofimmaturestagesuntilmidYJune，butincreasedftomlessthan2OpercoIonyduringJulytoapeak
OF56．8土S．d．36．8（n＝12）inlate September．Fe＼＼・lar＼′ae＼、・ere foundin coloniesin Tnid−November．
ThecocoonsofworkersfirstappearedinearlytomidJune．Themeannumberofcocoonsofworkers
increasedgreatlywithamaximumof75．4土S．d．32，3（n＝5）percolonyinlateAugustandthereafter
declinedrapidlybylateSeptember．Malecocoons rirstappearedinsomecoloniesduringearlytomid−
AugustandwerenotfoundinanyofthecoloniescollectedduringtheperiodfromlateAugusttoearly
SeptemberandreappearedafterlateSeptemberineverycolony．Newqueencocoonswerefirstobserved
inmidpSeptember．CocoonsofbothsexualswerefbunduntilearlyNovember．Themeannumberofmale
COCOOnSreaChedapeakof49．9土S．d．44．1（n＝7）ineariyOctoberandthatofnewqueencocoonsfluc−
tuated鉦om2．6i＝S．d．2．8（n＝7）inearlyNovemberto20．9よS，d．26．8（n＝7）inearlyOctober．Themax−
imum numberofmaleand newqueencocoonspresentpereolony＼、・aS127（A7519）and80（A6821）in
earlyOctober，reSpeCtively．
Comparati＼’ebioLogyorthc riveJapancsespecieso＝llCgenuS 拍spu（Hymenoptera、＼七spidac）
10l
V：simi〟ima：Themeannumberofeggsprcolonychangedfrom7．3土S．d．3，8（n＝3）inlateMayto
354・2土S・d・208・5（nニ9）inmid−October，andthereafterdeclinedduringNovember，thendisappeared
inearlyDecember，Themaximumnumberofeggspresentperco10nyWaSl，255inanest（S7010）collected
On September17th，1970．
Themeannumberofimmaturelarvaefiuctuatedapproximatelyequallytothatofeggsthroughoutnesting
Period．ThemaximumnumberofimmaturelarvalstagespercoIonywasl，218inthenestmentionedabove．
Themeannumberof5thinstarlarvaewasalsoapproxirnatelyequaltothoseofimmaturelarvalstages
untilearlyAugust，andtherea触rnuctuatedfrom22l．3j＝S．d．208．9（n＝8）inmid−Augustto127．0土
S・d・18・4（n＝3）inlateNovember、Vithamaximumof588．4士S．d．392．3（n＝10）inlateSeptember．The
maximumnumberof5thinstarlarvaepresentpercolonywasl，336inanest（S6902）collectedonOctober
4th，1969，
CocoonsfirstappearedinearlyJune．Themeannumberofworkercocoonsincreasedrapidlycom−
Paredwiththatoftheother Vh5PaSpeCies，andreachedamaximumof563．4土S．d．496．7（n＝10）inlate
Augustandthe陀a托erdecreasedto2．2土S．d．4．3（n＝12）inlateOctobeちthendisappearedbymid−Novemben
MalecocoonsfirstappearedinmidTAugustinafewcolonies，butwerenotfbundinanycoloniescollected
inlateAugust．Duringtheperiodfromearlytomid−September80utOf24coIoniescontainedmaleco−
COOnSandthepercentageoFtheco10nies、、・ithnlalecocoonsreached80町0（8／10）inlateSeptember．The
meannumberofmalecocoonswas628．0土S．d．532．1（n＝＝11）inear董yOctoberandincreasedto668．0
土S．d，449．4（n＝12）inlateOctober，andthereafterdecreasedrapidlyduringNovember．Themaximum
nunlberofmalecocoonspercolony＼、・aSl，743inaqueentessnes＝S7021）collecLedonOctober21st，1970．
NewqueencocoonswerefoundduringtheperiodfromlateAugusttoNovember．Themeannumberof
COCOOnSPrCOIonyremainedlessthan20untilmid−Septernberandthereafterincreasedrapidiyto511．6
土S．d．342．0（n＝12）inlateOctoberandthendeclinedduringNovember．Themaxirnumnumberofnew
queencocoonspercolonywasl，380inanest（S6606）collectedonOctober27th，1966，
3）AduIthornets：
V：trqpica：WorkersbegantoemergeinlateJulyandafewofthemweresti11presentinmid−October
（Fig．71），ThemeannumberofworkerspercolonywasO．4土S．d．0．9（n＝5）inJulyandincreasedto
S当字叫旨・0紙
LL
12．6土S．d．9．2（n＝12）inAugust，andthereafterdeclinedduringSeptember．Themaximumnumberof
●「■￣−、
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Fig．7l．Seasonalchangesinnumberofadults主nco夏oniesof掩叩atrOPEca．Opencircleandvcrticallineinp
dicatcmonthl〉′】11Can and st；lndard devialion．
102
Makoto MATSUURA
、VOrkerspresentpercolony、＼raS40inanest（T7303）collectedonAugust23rd，1973．Malesbegantoemerge
inlateAugust．InSeptemberthemeannumberofmalespercolonywas4．9土S．d．8．9（n＝13）witha
maximumof30percolony（T7104）．NewqueensalsobegantoemergeinlateAugust．Themeannumber
Ofnewqueenswasapproximatelyequaltothatrnalesthroughouttheemergenceperiod．Themaximum
numberofnewqueen percolony＼、∫aS32in anest（T7105）co”ected on September19th，1971．
Vぐ用bTO：Workersbegantoemergefrommid儀tOlateJune（Fig．72）．Themeannumberofworkers
increasedfromlO．3＝たS．d．6．7（n＝3）inJulyto162，4土S．d．74．2（n＝12）inSeptemberwithamaximum
Of426inacolony（C6802）co11ectedonAugust30th，1968．Itdecreasedrapidlyto52．2土S．d．583（n＝11）
inOctoberandworkersdisappearedinmostco10niesinearlyNovember．Thefirstmalesappearedfrom
mid−tOlateAugustinafewco10nies．Themeannumberormaleschanged rroml17j＝S・d・411（n＝7）
inAugustto51．5土S．d．130．1（n＝11）inOctober，Themaximumnumberofmaleswas434inaqueenless
nest（C7508）collectedonOctoberlst，1975．Newqueensfirstappearedinafewcoloniesfrommid−tO
lateSeptember．Themeannumberofne≠′queenSWaS38．9土S．d．71．2（n＝11）inOctober＼＼rithamaximum
of217inanest（C7405）collectedonOctober13th，1974．InNovembernewqueenswerenotfoundin
O−O
︶
anycolonies．
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−
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0。
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Fig．72．Seasonalchangesin numberoradultsincoloniesof FJt，3PL7Cmbro．OpencircleandverticaL Linc
iIldicate nlOnthlymean andstandard deviation．
r：ma（1darinia：Wbrkersbegantoemergefrommid−tOlateJune（Fig．73）．ThemeannumbeI・Of＼＼・Orkers
increasedfrom47，0土S．d．46．7（n＝2）inJulyto178．6土S．d．140．2（n＝4）inSeptemberwithamaximum
Of514inanest（M7202）co11ectedonSeptember19th，1972．Thereafterthenumbersdecreasedto91．1
土S．d．60．2（n＝16）in No＼′ember．
MalesfirstappearedinafewcoloniesfromlateAugusttoSeptember．Themeannumberofmalesin−
CreaSedFromll．9j＝S．d．30．2（n＝14）inSeptemberto74．1土S．d．97．0（n＝16）illNo＼′ember＼VheIlallthe
acti＼′eCOloniescontained males．Themaximumnumberofnlales present percolony＼＼・aS284inanest
（M6708）co11ected on November25th，1967．
NewqueenswerenotfbundinanycoIoniesuntilNovemberexceptforonenestwithonly8queens
ComparativebiologyofthefiveJapanesespeciesofthegenus侮pa（Hymenoptera，Ⅶspidae）
103
3
2
S山一＜芝
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。
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MAY JU N JUJ AuG S E P O C T N O＞ D E C
Fig．73．Seasonalchangesinnumberofadultsincoloniesof
挽卿ⅧW融加地Openeircleandverticalline
indicatc monthlymean and s【andard dc＼′iation．
2
0
〇
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8
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Fig・741Sea50naichangesinnumberofadult5inco始nieso仁撤御用皿撤（10−daymeans）．
（M7402）coilectedonOctober17th，1974・Therneannumberofnewqueenspercolonywas74．1土S．d，
97・0（n＝16）inNovemberwithamaximumof396inanest（M7403）co11ectedonNovember2nd，1974．
Adulthornetswerenotfbundincoloniesexcavatedin December．
Vana蝕：WbrkersbegantoemergefromlateJunetoearlyJuly（Fig．74）．Theirnumbersdidnotexceed
20percolonyuntillateJulyandthereafterincreasedrapidlyduringthetimen・Ommid血gusttoearly
SeptembenThemeannumberoぎworkerspercokny加ctuatedbetween弧7土S．d．24．？（n＝5）inlate
104
Makoto MATSUURA
Augustand54．9土S．d．52．5（n＝9）inmidAugust．Numbersofworkersdecunedsharplyaftermid−October，
and＼l・OrkeTSdisappearcdjnmjd−November．Thcmaxjmumnumberor＼＼′Orkerい＼′aS142inanest（A7513）
CO11ected on August20th，1975．
MaleswerefirstfoundinafewqueenlesscoIoniescollectedfromearlytomid∼August，andthereafter
disappeareduntilmid・September．Whenrnalesreappeared，themeannumberpercolonychangedfrom
8．4土S．d．18．2（rl＝5）irlrnid−SeptembeTtO46．8土S．d．28．3（n＝9）inearlyOctoberwithamaximum
Of84malesinanest（A75i9）collectedonOctober7th，1975．Theirnumbersthendecreasedsharplyt
2．1±S．d．3．8（n＝8）in early November．
Newqueensbegantoemergefromrnid−tOlateseptember，Themcannumberofthenewqueensper
COIorlyfluctuated＆ommid−OctobertoearlyNovemberbetweenlO．8i＝S．d．19．2（n＝9）inlateOctober
and12．1土S．d．23．3（n＝8）inearlyNovemberwithamaximumof77inanest（A7111）collectedonOcp
tober19th，197l．Adulthornetswereabsent fromcoIoniesbymiかNovember．
V：simiHima：Thefirstworkersbegantoappearfromrnid−tOlateJuneandthereafterthenumberof
WOrkersincreasedrapidly（Fig．75），COmparedtothatoftheotherspecies．Themeannumberofworkers
い﹂＜⊃ユー＞ココ〓一Jご．eこ
800
∧︶
∩）
〔）
0
MAY JUN 」UL AUG S E P O CT NOV DE C
Fig．75．Seasonalchangesinnumberofadultsinco王oniesof tepasEmi［／ima（10・daymeans）．
percoionyincreasedtoover120bylateJulyandattainedamaximumnumberof574．9d＝S．d．395．8（n＝ll）
percolonyinlateSeptember（Fig．75）．DuringOctobertheirnumbersfluctuatedbetween442，6土S．d．
356．5and501．7土S，d．489．7，anddeclinedsharplyduringNovember．Theproportionofeolonieswith
WOrkersover400was54，5甲0（6／ll）incoIoniescollectedinlateOctober；butitdecreasedto25．0（殉（2／8）
in eal■ly November．Workers werestillfoundin somecolonies fromlate No＼′ember to early December
anddisappearedatmid・December．Themaximumnumberofworkerspresentcolonywasl，384irtanest
（S6603）collected on October5th，1966．
Maleswerefirstobservedintwoqueenlessco10niesinearlySeptemberandthereafterappearedboth
inqueenrightandqueenlesscolonies．ThemeannumberofmaiespercoionylnCreaSedfrom31．5士：S．d．
77．1（n＝13）inearlySeptemberto357．4±S．d．301．6（n＝8）inearlyNovember，nuCtualingatlevelsof
OVer300percoIonyn・Ommid−OctobertoearlyNovember，andthereafterdeclinedrapidiyto6．4士S．d．
9．O（n＝5）inlateNovernber，thoughmanymorewerefbundinafewcolonieseouectedinearlyDecember．
Themaximumnumberofmalespresentpercolonywasi，814inaqueenlesscolony（S7315）coilectedon
October5th，1973，Whichhadneverproducedanynewqueens，and661inaqueenrightcolony（S66O5）
COllected on October27th，1966．
NeⅥ′queenSbegantoemergeinafewcoloniesfromearlytomid−September，andthereaftergradua11y
increaseduntilmid・November．Themeannumberofnewqueenspercoionychanged丘om31．5土S．d．
77．1（n＝13）inearlySeptemberto180．6士S．d．112．5（n＝5）inmid−November，nuCtuatingatabout135
duringmid−Octobertomid・November．Thereafterthenumbersdeclinedrapidlyto2．3よS．d．4．0（n：＝3）
Comparativebiolog）・OrthcfiveJapanesespeciesorthegenuslセ∫PO（Hymenoptera．Vespidae）
105
inearIyDecember・Themaximumnumberornewqueenspresentpercolonywas350inanest（S6811）
COllected on November3rd，1968．
3．SeasonaldevelopmentofaslnglecoIonyof掩軍atrqPica
Itisverydiffieulttofollowthedevelopmentofindividualcoloniesof物aduringtheiractiveperiod
fromnestfoundationtodisintegration，becauseviewsoftheinnernestareinterruptedbyathickenvelope，
espcciallyatthelaterstageofnesting．However，in Vtropicatheenvelopeisincompleteandthelowestcomb
isalwaysexposedwithinthenestcavity・ThisfacilitatesdirectobservationofpopulationsinsideacoIony
withotltdisturbingittoomuch・ChangesinthecoIonydevelopmentandpopulationstructureofaslngle
nest（T7401）ofレ：tropicaareshownin Figs．76and77throughouttheacti＼′eSeaSOn．
山○
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31 10
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Fig・76・Seasonalchangeofbroodpopulationsinalセspo（rOPicacolony（T74011．Årrovvsdeno【eSthcda）rOf
the点rsモemergenceoreaehcaste、
l）Cellconstruction
王nthiscoIonythequeenbeganto foundhernestonJune9th，1974andbuilt42cellsaloneuntilthe
time＼VhentherirstworkersemergedonJul〉′14th．1ntheperiodfromnestinitiationtoworkeremergencc
themeannumberofcellsconstructedperdaywasl、1土S．d．l．3（range：0∼6）．A餌rworkeremergence
thenumberofce11sincreasedrapidlyandasecondcombwasbuiltonJuly2OthandathirdoneonAugust
7th，allthesemainlyconstructedbyworkers．ThenestreacheditsmaximumonAugust2lst；atthetime
thenestwascomposedof268ceuswithin3combs．Themeannumberofcellsconstructedperdaywas
4．9土S．d．3．4（range：0∼15）duringtheperiodfromworkeremergencetofinalcellconstruction．The
numberofemptycellswasveryftwthroughoutthesolitaryperiodandrarelyexceeded6．0鴨ofthetotal
numberofcellsuntiltheemergenceofsexuals，becauseemptyceIIseithernewlyconstructedorafter
emergenceweregenerallysuppliedwithaneggassoonastheywerevacant．Thereafteちtheemptyceus
increasedrapidlyduetothecessationofovipositionandcannibalismofbroodstages．
2）lmmaturestages
Asthequeenlaideggsonlyinnewlyconstructedce11sduringthesolitaryperiod，therateofovlpOSition
WaSVerySimiiartotbatofcellconstruction．Shelaidanaverageofl．4e名題5土5．d．1−8（range：0∼4）per
dayfbrthefirst35daysfromnestinitiationtothefirstappearanceofworkers．Afterworkeremergence
106
Makoto MATSUUR∧
Fig．77．Dailychangc ofaduIt populationsin alセ3Pa（rOPicacolon〉・（T7401）．
theovIPOSitionrateincreasedrapidlyandthehighestratereached14eggsperdayonJuly27thand28th．
Thehighesttotalnumberofeggswas76，Observed15and16daysafterworkeremergence．Thereafter，
thetotalnumberofeggsdecreasedwithoutrecovery，thoughitvariedbetween34and48duringAugust
anddeclinedrapidlyinearlySeptember．ThequeenstoppedovipositingonSeptcmber13thanddiedon
September20th，andalleggsdisappearedonSeptember14th．
lnthelarvalstages，lstto4thinstarlarvaereeachedtheirpeakintotalnurnberinearlyAugustwith
amaximumof731arvaeonÅugust5th，anditresultedinariseinnumberof5thinstarlarvaea魚erabout
One＼Veek＼＼・hen theyoccupied nearly75甲00fallthelar＼′aepreSent．
ThenumberofcocoonsreachedapeakinlateAugustwhentheyoccupiednearly52．5％ofa11the
brood with a maximum of137cocoons perdayonAugust25th．
Allthebroodstagesdeereasedrapidlywiththecommencementoftheernergenceofsexuals．Eggsand
immaturelarvaeoflstto3rdinstarsfirstdisappearedonSeptember14th，and4thinstaronthefo110Wing
day．CocoonsdisappearedonSeptember21st，butthe5thinstarlarvaeremaineduntilSeptember24th．
3）Adulthornets
Thefirstworkeremerged36daysaftercolonyfbundation．Theincreaseinthenumberofworkerswas
VerySlowduringtheearlynestingperiod，andthetotalnumberofworkerswaslessthanlOforthefirst
18daysafterfirstemergence．Emergenceofworkerslastedfor44daysandstoppedonAugust28th．The
numberof＼VOrkersreachedamaximumof600nAugust26thand27th，justbeforeemergenceofsexuals，
andthereafterdeciined．ActivityworkerwasobservedasiateasOctober王4th．Thetotainumberofworkers
producedinthiscoIonywas71，andtherateofemergencewasl．7土S．d．1．2（n＝44）workersperday
withamaximumof6workersperday．
Comparaモivebio10gyOrthenYeJapanese叩eCiesofthegenus物♂（Hymenoptera，職印idae）
107
Thef3rstmalesemerged67days，andthefirstnewqueensemerged74daysaftercolonyfbundation．A11
malesemergdin25daysandamnewqueensin18days；thelastemergenceorbothsexudswasonSeptember
19th・Thetotalnumberof＄eXualsproducedinthiscolonywas6lmalesand75ncwqueens．Anexplosive
emergenceoccurredinthefirsthalfoftheemergenceperiodwithanincreaseto18and20queensperday
OnSeptemberlstand2nd・Bothsexualsstayedfbrabouttwoweeksinthenest：malesbegantodepart
OnandafterSeptemberlst，neWqueenSfromSeptemberlOth・AllhadleftthenestbylateSeptember．
4．Production ofsexuals
l）Numberofscxualsprodueedpercolony
Thenewqueensandrnalesareproducedinthelaterpartofco10nydeYelopmentwhenthenumberof
WOrkersreachesapeak．職ble39showsthespeciricdi胱柁nCeintotalnumberofsexualsinautumnnests
Offive物aspecies，baseduponbothadulthornetspresentinthenestandcocoonsofeachsex，Which
籠ble39・Thenumberofsexualsproducedpercolonyinfive掩spaspecies．
No・OrneWqueモn＄Pre5en【 No．ormalespre5enl
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9，323
FiglLreSin paTCllthescs dcnole S．d．（土）．●No．orncsts obser＼・ed．
COuldbeeasilydistinguishedfromoneanotherbytIlethicknessandlengthofcellcapplngaSrepOrted
bySpRADBERY（1971）andMacDoNÅLDetal．（1974）知r掩岬〟ぬspecies．Themaximumnumberofsexuals
PreSentVariedfrom56newqueensand27malesina f（trppicanest（T7303）tol，692newqueensand
l，123malesina VsimillimGneSt（S6605）andthenumberofsexualsproducedperco主onyisrankedas
頭痛肋肌用㌧傲紬弛め痴1＝肌勅℃＞8胱ぬ＞′J叩おα．
itseemslikelythattheproductionofsexuais，eSpeCiallynewqueens，dependsgreatlyuponthecolony
Sizenotonlyamongdi胎rentspeciesbutalsoamongconspeciriccolonies，thou如correlationbetween
numberofcel15andthatofnewqueensproducedinthesamespeciesisnotclearexceptぬr打ぷ∼椚′／肋α
（職ble39）．Inthecaseorシ∴扇〝痛里れ柑tbereseemstobeastrongcorrelationbetweenthenumberofnew
queensproducedwiththatofceusconstructed，eSPeCia11yintheneststakeninOctot）er（Fig．78）．The
108
Makoto Mノ汀SUURA
■U
0
O
2
八Sト﹂⊃n︽Sヨ乱S喜OUOU︶S宏山⊃8透冨山〇．OZ
2000 ▲000 dOOO 80001000012000
NO．OF CELLS PE尺 NEST
Fig．78．Correlationofnumberofcellswithnumberofnewqueenspresentpercolon〉IOflセ5PaSimi／／imotaken
duringOctober．
1argesuccessfulcoIoniesexceedingca．8，000cellshadusuallymorethanl，OOOnewqueenspresentaswe11
asmales，＼＼・hileapoorcolonysuchasS7317with949ce11sinOctoberhadonly16queens，thoughsorne
mayhavebeenproducedinthecourseofcolonydevelopment．
Thismaysuggestthatthenumberofcellsinareproductivelysuccessfulco10nyOf Vsimi／Iimaneeds
toreachacolonyleve10fatleastl，000ceLIs，andontycoloniesattainingsufficientsizeha＼′etheability
toproducenewqueens．
Comparingthisminimumsizewiththeotherspecies，thenumberofcellsproducedbefbreandduring
thenewqueenrearingperiodisestimatedonthebasisof7hble39（thoughdataoneachspeciesarenot
Su汀icientinnumber）as如low：巨沼α〃（ねrf〃料C臥800cells；Hc／V∂明C乱300ce11s；巨α〃α梅200ce11s
and レ：trppicalCa．100cells．
Ontheotherhand，SOmeOfthenestswhichhavelosttheirinitialqueenbeforebeginnlngqueenprO−
duction reach oroften exceed such a numberofcells，but produceonlymalehornets．
2）Larva：WOrkerratio
InadditiontotheadequatecolonysIZeinthenurnberofcellsineachspecies，itisrecognizedthatthere
existseveralotherfactorswhichpromotenewqueenproduction．Thechangeinthelarva：WOrkerratio
isoneofthemostimportantfactorswhichfavourstheproductionofnewqueensasreportedinbumble
bees（R靂CHÅRDSandRICHÅRDS1946，CuMBER1949）andinothersocialwasps（RICHARDSandRICHARDS
1951，SpRAD8ERY19乃，ARCHER1972）．
Fig．79showstheseasonalchangeofiarva：WOrkerratioincoloniesoffive掩軍aSpeCies・Theratio
remainedgenera11yhighduringtheperiodpriortotheproductionofnewqueensinallspecies・Theratios
becamemuchlowerbeforetheperiodwhencoloniesbegantoproducequeenbrood，andduringtheperiod
whencolonieshadrearedqueenbroodtothecocoonstage，theratioapproachedlessthan2．0．Thenthe
ratiobecameirregularasthenewqueensandmalesbegantoemergeinthenestL
Ontheotherhand，itwouldappearthatthechangeinlarva：WOrkerratioisconcernedbasicallywith
thenumberofworkerspresentineachspecies．Fig．80showsthatthelarva：WOrkerratioisrelatedtothe
numberofworkerspresentincoloniesof Vstmil［imaand Vana［isbefbretheperiodoftheemergence
Comparativebio10g）rOfthefiveJapanesespeciesofthegenus咋5Pa（Hymenoptera，Vespidae）
109
8
志謹皇＼V姦3
ムー
一句
つ↑
占Ⅹ
∩︶ノ0
■q
2
0
8
る
d
2
0
10
8
d
d
2
0
Fig．79．Seasonalchangesoflarva／workerratioincoIoniesoffive物aspecies．Opencircles：COIonicswith
Onlyworkel●brood；SOlidcircles：COlorlie5Wi抽newqueenspresent．
6
8
4
2
0
誓〓壬￥＼V、⊥く一．︹∴
6
∝山寓∝￠貰、ペ＞∝ぺJ．〇麗
8
4
6
4
2
2
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6
0
150
iJO．∧DULTlイORKE柁S
4
2
lOO
V＄irlli＝m8
∵∴・・ら・−・・
200
∠‡00
600
800 1000
‖0．A祝し7一書ORKERS
Fig．80．Comparisonoflarva／workerratiosinrelationtot王1enumbersofworkerspresentincoloniesof V
Simil／iITl〟（lo、＼・Cr）andl／：urla／i∫（upper）bcrorclhcperiodorthce）11ergCnCeOrne＼＼・queC）1S．OpellCircIcs：
newqueerlCOCOOnSpreSe11t；SO】id circles：WOrkerbroodonly．
Makoto MATSUURÅ
8
6
∝山叉∝0王＼く＞∝5．〇〓
4
0
300
・100
500
NO．∧DULT＼ヾORIこERS
6
∝山虫∝0‡＼く＞∝5．〇芝
8
4
100
200
300
480
500
NO．A王）ULT WORKERS
Fig・8l・Lar＼■a／＼l■Orkerratiosorthetwo柁5PaSPeCies、lてcrabro（upper）and［てmqndariltai（lo≠・er）．Opencircles：
newqueencocoonspresent；SOlidcircles：WOrkerbroodonly．
Ofnewqueens．DespitetheremarkabledifferenceincoIonysizebetweenthetwospeciesthereexistsa
COnSplCuOuSSimilarityinlarva：WOrkerratios；theratiosinbothspeciesareapproximatelyequalandthe
Valuesfbrthequeenrearlngperiodvariedonlybetween2．Oandl．0．In Vsimi］［imasuchvaluesinthe
ratiosoflarva：WOrkerwerelikelytobeconstantamongcolonieswithmorethanabout300workers，and
thevaluesinthecolonieswith50∼300workersvariedalmostentirelybetween3．Oand2．0，andthos
inthecolonieswithlessthan50workerswereverymuchhigher．Ontheotherhand，inthecaseof V
analistheratioftubelow2．Owhenthecolonyhadmorethanca．40workers，anditincreasedrapidly
in the colonies withless than ca．20＼VOrkers．
Fig．81showsthelarva：WOrkerratiosofcoloniesofVcTqbtvand VmandarinjLLWhichareverysimilar
incolonysize．Inthesespeciesvaluesforthelarva：WOrkerratioalsovariedapproximatelybetween2．O
andl．OincoIonieswithmorethanca．10Oworkersduringthequeenrearlngperiod．
Judgingfromthesedata，itseernslikelythatareductioninthelarva：WOrkerratiotoapointbelow
2．OwithanincreaseofthewoTkerpopulationwhichmaintainsacertainbalanceineachspeciesisaslgnal
fbrqueenproduction，thoughworkerpopulationforrearlngqueenbrooddiffersconsiderablyamong
掩5pαSpeCies．
5．Nest sur＼うval
l）Survivorshipofcolonies
InI句卯SpeCiesitisdifficulttoobtainreasonablya亡CurateeStimatesofthesurvivorshipofcolonicsfrom
thetimeofnestinitiationbecausethedensityofnestsinthefieldisgenerauymuchlowerthanthose
Comparativebiolog）・Ofthe fiveJapanesespeciesorthegenuslセ5Pa（H〉・menOptera，Vespidae）
111
OfotherJapanesesocial＼VaSPSSuCh as Po／Lf（esand L，＞spzL／aspecies．
Onlyasmallamountofdataareavailableonthenestof VanalLs．From1964to1975，Ilocated59
nestsofthisspeciesandobservedthemfromthesolitaryperiod：28（47．5甲b）losttheirqueensbeforeworker
ernergence，8（13．6甲b）diedoutduringthecooperativeperiod，3（5．1％）failedtoproducesexualsinspite
Oftheirworkerproduction，2（3．3甲b）weredestroyedby Vmandariniawhentheymatured，andonly18
（30．5勒）survivedtoproducenewqueens．ThesefiguresshowthatinIくanaIi5alargenumberofcolonies
Perished duringthesolitaryperiod mainly because ofthelossoftheirqueen．
rrhble40indicatesthenumberofunsuccessfulnestsinthefive噸aspeciesbaseduponmyobserva−
tionandtheanalysisofnestcontentsatthetimewhenthenestswerefoundinthefield．Thedatafrom
SuChnestsglVeSOmeinformationwithwhichtoestirnatethesurvivorshipo‖句卯COlonies，andvarious
factorscontributingtocolonyterminationareglVeninrrhble41．
Thble40．Thenumber ofunsuccessfulnests at each de＼・elopmentalperiodin rivelセ5PaSPeCies．
Solitary＊
Species
1
7
レ：∫／〝！J／／山川
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l1什叫如m
4
5
3
6
7
（10．0）（20．0）（23．3）
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7
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29
1btal
Cooperative Polyethic Reproductive
58
（8．2）（11．8）（23．7）
6
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9
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7
（14．3）
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（40−7）
4
37
（軋さ）
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245
（20．4）
＊Ⅰ：befbrehatchingoflarvae，ⅠⅠ：fromhatchingoffir＄tlarvaetosplnningoffirstcocoon，iII：thereaftertitl
抽eemergenceortherirstworker．Parenthesized軸uressbow甲o ratioineachperiod．
In Vana／ねterminationofthecoIonywasobservedmostfrequentlybefbreworkerernergence，Which
included51．9甲oofallunsuccessfulnestssofarobserved．Asecondaryperiodofhighmortarity．thesuc−
cessi＼′eCOOPerativeperiod，reSultedinalossof2l．6qo．Thesecasesseemtobeapproximatelysimi1arto
thepreviousdata．Suchgreatcolonymortalityin Vana姑seemstobemainiycausedbythelossofthe
queenduringherforaglngPeriodsfromthesolitarytothecooperativeperiod，and49．0％outofunsuc−
CeSSfulnestslosttheirqueeneitherbeforeorsoonafterfirstworkeremergence．
IncoloniesofVsimillbnawhichoftenchangetheirnestingsiteafterworkerernergence抒omtheoriginal
c10Sedplacetoanopenplace，32．6％ofa11theunsuccessfulnestsdiedoutduringthesolitaryperiod
befbremovingfromtheiror唱1nalnestingsite．Thehighestrateofcolonycessation，40．8％ofalltheun−
successfulnests，WaSObtainedduringtheperiodaftertheemergenceofsexuals．Thishighermortality
insuchmaturedcolonieswasmostlybroughtaboutbyattacksof V mat！darinia．
In VcTUbrv53．3勒oftheunsuccessfulnestsfailedduringthesolitaryperiod，and30．O甲binthecooperative
period．Inbothcasescolonyterminationwasmostlycausedbythelossofthequeen，andinthelatter
case，Of9nestsperishingduringthecooperativeperiod，threewerecausedbytheiossofthequeenwhich
112
Makoto MATSUU】iA
’rhble4l．Fac【OrS亡Ontributingtocolon），terminationin fi＼′Clセspaspecies．
ロ′用地
Sp亡Cie5
Period ●
∫f川〟〟／J和
A＋B
／机川血√血血
ぐ川上〝0
A＋B
A＋B
1blal
／叩小川
A十B
A＋B
A＋B
A B 勒 A I】甲o A B t穐 A B （恥 A B （穐 A 8 甲0
Lossorqueen
50 （〉 50 3 5 8 9 0 9 8 0 8 18 2 20 88 7 95
（49．0）
Naturalenemies
AttacL by
（16．3）
0 0
O ll ll O O O
honeybuヱZards
（敗8）
AtlaCkby挽鱒α
（30，8）
（30朋
0
0 0
0
（0．0）
（0・0）
0 0
0
（54．1）
（38．9）
岬．0）
四．5〉
0 11 11
（0．0）
0 6 6 0 17 17 0 3 3 0 0 0 0 1 1 0 27 27
J〃〃肌ねr′〃由
（5．9）
（34．7）
（10．0）
（0．0）
（2．7）
（11．1）
Stylopiza血nby O O O O O O l O l l O 1 0 0 0 2 0 2
〟ビ／JO∫ぐ用わ′0／＝∫
Parasitism by
lepidoptemn
Hunlallinl⊂rr亡†e11CC
De5truClionbyman
Erkc10rin5eCtト
Cidespray
Er托c10fland
reclamation work
Climatic fもctors
Strongwindwilh 7 5
heavypreclpiほtion
Unknowrl払ctor5
（0．0）
（0．0）
（3．3）
（3．8）
（0．0）
（0．8）
（0．0）
（0．0）
（0．0）
（0．0）
（Z．7）
（0．4）
O O O O O O O O O O O O O l ユ 0 1 1
3 3
6 1 3
（5．9）
4 2
6
（5．9）
2 1 3
（8．2）
2 1 3
4 0
0 10 10 0 0
（38．4）
（肌0）
4
（6．り
0 0
0
0 2
0 （；17 23
（0．0）
（9．4）
2 10 5 15
（13．3）
（0．0）
（5．4）
（6．Ⅰ）
（0．0）
（0，0）
（0．0）
（0．4）
O O O O O O O O O O l 1 0 0 0 0 1 1
（8．0）
（0．0）
12 0 0 0 0 0 0 1 2 3 1 0 1 9 7 柑
（11．8）
（0．0）
（0．0）
11 0 11 】る 1 17 9 i iO 3 1 4
（氾8）
てbla1
4
（34．7）
（33．3）
（11．5）
（2．7）
（6．5）
7 5 12 4る 8 54
（】5．4）
（32．4）
（22．0）
75 ヱ7 102 2之 27 49 25 5 30 王3 i4 27 26 】1 37 1る1 84 245
＊Å：perio雨rroIⅥSOlitarytく）COOperativeperiod，8：periods抒ompolyethictoreproductiveperiod．Pare】1thesizedr短ures5how勒ratio
throughouもA＋B．
COuldnotlocatethenewnestlngSiteinaconceaiedplace，aSthisspeciesalsotendstochangeitsnesting
Sitewhenthenestspacebecomestoonarrowforcombbuilding．
InthecaseofVmanddrin如hediscoveryofnestsisverydifficult，eSpeCiallyaモtheearlieststage，because
Ofitsexclusivesubterraneannidification．TheperiodofgreatestcoIonyperish重ng，reSultingin40．8％of
autheunsuccessfulnests，WaSWhenthenestrnaturedandproducedsexuais．Aftertheemergenceofsex−
uals100utOfllneststhathadperishedhadbeendestroyedbybeekeeperswhosoughtthenestsbecause
thisspecie5CauSeSSeriousdamaさetOhoneybeecolonies．
1nI′．（rOPica40．5町00ralltheunsuccessfulnests raileddul■ingthesoli【aryperiodand29．7勒inlhe
COOPerativeperiod．Of37nests，18diedoutduetothedisappearanceofthequeen，PrObablydtlring
roraglmg．
Judgingfromthesedata，ingerieral，thedemiseof掩軍aCOIoniesseemstooccurvery鉦equentlydur−
1ngthesolitaryperiodandthecooperativeperiodwhenthequeenstillcontinuestofbrageoutsidethe
nests，tllOughthereexistsomespeciricdi胎rences．
2）Morは1ity払ctors
7hbIe41alsoshowsmortalityfactorsafftctinghornetsmainlyatthenestleveIratherthanattheindividual
Comparativebiologyofthefi、・eJapanesespeciesofthegenuslセ5Pa（Hymenoptera，Vtspidae）
113
1evel・These払ctorsopemteon物αCOloniesinasimilarwaytothe“a110rnOne”action陀pOrtedぬrag−
gregationsoftheimmaturestagesofastinkbug（HoKYOandKrRITANZ1964）．Fivemortalityfactorsare
recognized：（1）disappearanceofqueen，（2）naturalenemies，（3）humaninterftrence，（4）climatic払ctorsand
（5）unknown causes．
（l）Disappearanceofqueen
From職ble4いtisestimatedthatthemaincauseofcoIonydemisein挽軍aisattributabletothedisap−
PearanCeOfthequeen，particularlyatanearlystageofnesting・Among245nestsofthe蔦ve挽甲aSpeCies
exarnined，theseorphanednestsrepresented38・9％ofthetotalunsuccessfulnests，thoughtheratioofnests
tbatlosttheirqueenra喝ed什omlO▲0％orunsuccess蝕1nestsor巨c′Wわ和tO54．1勒orthoseorr仇p加．
Theremaybesomephysicalorbioticfactorswhichcausethedisappearanceoffbundingqueens，SuChas
naturalenemies，Climaticconditions，aVailabilityoffbod，humaninterference，Physiologicaldeath，failure
todiscovernewnestsitesinnestrelocationandevacuationasreportedinFb加esspeCies（YosHlXAWA1954）．
IobservedthattwofoundingqueensofVcTUbtvwhichhadmis＄edtheirnewnestsanddisappeared，and
OneOfthemhappenedtobekilledbythequeenandworkersofadi穐rentnestbecauseshehadmistaken
itfbrherownnestsite（MATSUURA1970）．However；itisgenerauyimpossibletodeterminethecauseofthe
disappearanceofthequeensintheirfbraglngtripsbydirectobservation．
Thefateoftheseorphanednestsmaybed11etOOneOfthefbllowing：（l）endofthecoIonycausedby
deathofthelarvaeandnewlyemergedworkersinthesolitaryperiod，（2）gradualdisintegrationofthecoト
OnyCauSedbysuccessivedisappearanceoftheworkersinthecooperativeperiod，（3）productionofmales
bylaying workers duringandafterthecooperativeperiod．Incaseof（3）theovariallydeveloped but
u11inseminated＼VOrkersproduceonlymales・grandsonsofthelostqueen．Thesemalesma）・beabletoafftct
theproductivityofthenextgenerationbyinseminatingsomenewqueenswhentheyappearinautumn．
Thenumberofsuchmalesproducedinorphanednestsiso鮎nmuchmorethanthatofnormalnestsat
thecorresponding period．
（2）Naturalenemies
TherearefournaturalenemieswhichareregardedasmortalityfactorsoperatingatthecoIonylevel，nOt
attheindivid11allevel．Theseare“colonypredators”suchasthecongenericspecies，Vmanddriniaandthe
honeybuヱZard，Fbm豆ap［vonJS．L．
Vmandariniaattacksnestsofothercongenericspecies．OncediscoverlnganeSt，SeVeraldozensof V
mandarb7iaworkersvisititandacolonyisahnostcompletelydestroyedina鎚wdays．Theatt＆Ckisseasonally
Welldelimitedandcatastrophicdamagealwaysoccurs鉦ommid血gusttoearlyNovernbeちWhenthenest
reachtheirmaximumsizewithsexuals．ThisstrongpredationbyIてmaf2darin由causedthedeathofthe
highestproportionofalltheunsuccessfulnestsatthematurestage．Itisnoteworthythatthepercentage
Or粁ぶわ7∼〝f椚〃inparticularishi抽erthanthatof弧yOtherspecie5eXCe由払r挽甲〟ぬノ払γねぢば（取ble
20）．Itseemsukelythat Vsimillimaisdestroyedftequentlybecausetheirnestsareabundantinthedistrict
SurVeyed；alsotheirnestsarebuiltintheopenandinconsplCuOuSplacessuchasundertheeavesofbuildings．
ThecolonysIZemayalsoplayasignificantroleintheprefbrenceofVmandarinidfbrlてsimillimabecause
thenestoftheiatterismuchiargerthanthatofanyothercongenericspecies．
Thehoneybuzzard，勧isqpLVOruちisknownasapredatorof掩甲uJacoIoniesinEuropeancountries
（Sc＝REMMER1962，TRÅP−LJND1962，EDWARDS1980）．Thisbtlエardisnotverycommonintheregionssurveyed，
anditsattackon挽5PaSpeCieswaslimitedtothecoIoniesofVanalis．Of27nestsin VanaHswhichdied
Outduringandafterthepolyethicperiod，atleastllwereconsideredtohavebeendestroyedbythisbt暮Z−
Zard．AtanestofthisbuzzardIoftenobservedthatthebirdsbroughtpartsofacombintothenest，though
itwasnotdeterminedwhetherthebuzzardhadpreyedonadulthornetsornot．Oncethebuzzarddiscovers
hornets’nest，itkeep5VisitingitandplunderstherernaimngcombuntilthecoIonyiscompletelybroken
up．Anestattackedbythebuzzardcouldbeeasilydistinguishedbythelackofcombsinsidetheenvelope．
ItisnotcertainthatthebuzzardhasaspeCificpreferencefbrthenestof VanaHs；ratherthisspecies
maybeutilizedbecauseoftherelativelymildnatureofitscounteトattaekswhendisturbed，althoughIhave
neverseenanactualattack．Theexclusivelyaerialnests，tOO，areprObablyeasilydiscoveredbythebuzzard．
114
Makoto MATSUURÅ
Fig．82．Stylopizedworkersor柁岬αぐ用わ用Omtheenvelope．
Fig．83．Nest or停spa（rOPicq attacked by a
lepidopteran，たy用胎柁gαJね．
Vsimillimaisoneofthecornmonestspecies，aSalreadystated，btltthebuzzardmaykeepawayfromits
nestsbecauseofitshugecolonysizeandresultantseverecounter−attaCksbymanymoreworkersthanthe
case of 析α〃α撤
Asfrepsipteran，XeTlOSm7brvn豆Pierce，is occasionally foundasaparasite、、′ithinadult hornets（Fig．
82）．Thestylopizedhornetsspendmostoftheirtirnerestingonthecombwithinthenestwithoutdoing
anytasks，particularlyextranidalactivities，anda触rabouttwoweekstheyleavebutdonotreturntotheir
nest．Thelevelofparasitisrnofstylopizedhornetsisusuallyverylowinmaturecolonies：Ifoundonlyl，3甲b
Ofstylopized物aworkersinatotalof245，352adultsexamined，andtheefftctofastrepsipteranispro−
bablynegligibleinregulating停5PaCOlonies．Intheearl〉′StageOfnesting，ho＼、′eYer，theparasitismoften
reachedahighlevel．′RvocolonieseachofVmar7dwiniaand VctvbrDWeredestroyedbecauseautheworkers
emergmginthecooperativeperiodweresty10Pizedby．）おnoscrvb7Vn由andtheyle丘theirnestwithout
托turnlng．
1ha＼′eneVerfoundレhspanestscontaininglepidopterouslar、・aeduringtheiractiveseason，eXCePtingthe
OnereCOrdshownin′払ble41．OnthisoccasionitwasobservedthatseveraldozensofthelarvaeofPyru胎
regaIisBoisduvalinthece11sofIイtrqpicahadbeen feedingonthelarvaeandpupaein September1970
andhaddestroyedbothbroodandthenestfabric（Fig．83），despitethepresenceofseveralworkerhornets．
（3）Humaninterftrence
InJapan物ahornetsaregenerauyconsideredapesttobe飴ared，Withextremenuisancevalueandpoten−
tialhealthhazardcausedbystinglng．Thisattitudeexistsamongmostpeopleinal‖ocalitieswhereI句卯
SpeCiesoccur，andsomeofthemattempttodestroytheirnestswhenevertheyfindthem．7bmostJapanese
apiarists，Ontheotherhand，VmandariniaisaseriouspestcauslngCataStrOphicdamagetohoneybcecoト
Onies．Sotheysearchfbrthenestsofthisspeciesbyfbllowingthefiightpathofworkerhornets．Thedestruc−
tionsofnestsbymanplaysamoreimportan【partinco10nymOrta悠tyof VmandaT．iniacomparedtothe
Otherfourspecies，anddestructionbyaplaristsaccountedfbr38．4勒ofa11theunsuccessfulnests：mOSt
Ofthemwerekilledinthelaterstageofnesting．
Awidevarietyofinsecticidesseemstobeextremelytoxictoboth挽spalarvaeandadults．Some，SuCh
asearbaryl，dimethoate，diazinon，etCりWhichareadoptedinagrlCulturalsprayprogramsfbrpestcontrol
inorchards，CauSeSignificantmortalityifsprayedneartheirnests．Manyforaglnghornets，bothqueens
andworkers，diewhenpoISOneddirectlywithsuchinsecticidesinthefield．Someoftherncontactresidues
inthefieldandcarrycontaminatedwaterandfoodbacktotheirnest．Thesecausethedeathofnewly
Comparativet）iologyofthenveJapanesespeciesofthegenus物a（Hymenoptera，Vtspidae）
115
emergedhornetsas、、・ellasforagersthroughtrophallaxis．Suchhornetsmo＼′eaboutonandneartheground
bytheirnestbutareunabletoflyandtaketwotothreedaystodie．Broodismainlyki11edbychi11ing
OrStarVationduetothelackoffeedinghornets・Thele、′elofaccidentalkillingofcoloniesthroughthe
useofinsecticidesisestimatedat6．1甲bofthetotalofthefivespecies，butsomeofthecausesofdisap−
pearanceofqueensin fbraglngmayalso beattributabletotheeffectofinsecticides．
（4）Climatic factors
Weatherconditionsasdirectmortalityfhctorsappearstobeunimportantto物acoloniesundernormal
COnditions．However，StrOngWindwithheavyprecipitationoftenblewoff VanalLsnestswhichwereat−
tachedtotreesintheearlystagesofcolonygrowth．Ingeneral，neStSOf Vana胎intheearlystagesare
muchmoresusceptiblethannestsinthelaterstagestotheactionofstormsbecausetheformeraremore
fragileinstructurethanthelatterwhicharerigidlyconnectedwithtwigsaroundthenest．InfactIhave
neverfoundexamplesoflくa77alLsnestswhichweredamagedordestroyedbystormortyphooninthe
laterstageofnestingafterSeptember，Simi1arlylnaerialnestsofVsimillimG．Forspeciesofsubterranean
OrClosednidificationsuchasVmanddriniaand VITWicaheavyrainfalloccasionauycausedthedestruction
Oftheirnest，thoughaたwnestsofVmandat血iaandIてl叩tCaWeredestroyedasadirectresultof蔦00ding．
（5）UnkrlOWn払ctors：
ThelastlineofThble41show5thatunknown fhctorsaecountfbrhighproportionsofterminatednests
Of掩軍aSpeCiesthroughouttheiractiveseasons，thoughapartofthemmaybeattributabletooneof
the ractors nlentioned above．
6．Annualfluctuationsin thenumberofIセ5PqCOlonies
Table42showsthenumberofmature掩申aCOloniescollectedduringa12−yearperiodinandnearKibi．
1bble42．Fluctuationsin【hcnurnberofmaturcnestsin fi＼・■e L・を5PaSPeCieseollectedinandnear
Kibi，1964−1975．
Year
∫∼椚f′J血Ⅳ C相わ相
凸7
2
9
3
5
2
ハブ
エリ
Q′
3
7
′〇
l
Q／
﹁′− つ︼
つ︼ 3 4 5
57
・・1 7 3 5 つJ
0 1
7 7 フ 7 7 ︼7・
0ノ凸7 n7 Qノ
n7
′0
0ノ
3
八U
0′
‘U
00
‘U
0ノ
7
一エU
Oノ
′0
′〇
︵y
166
119
46
35
‘V 3 3 3 4 ‘V 4 ′0 8 ′0 ■1 3
2 3 つー 4 4 ■・1 3 4 4 4 つー．4
O
gJ
′0
Qノ
▲HV
ハU
Qノ
′0
Aて
Total
Tbtal
α乃〃′ね椚α〃（ね′f君ぬ r叩加
423
つー 1
Thesedatacannotbeuscdtodirectlycornpareinterspecificdifferencesinnestnumbersatthestudyarea，
n7
butthefiguresmayprovidearelativeindextomonitorinteトSpeCificrelativeabundances．
Thehigherrankofboth Vsimillimaand Vana胎mayrenecttheiraerialnestswhichareeasilydiscovered，
＼Vhilethelo＼＼∫erabundaneeoftheotherthreespeciesma），berelatedtotheirconcealednidifieation．ThereJative
1
abundanceamongthesefivespeciesestimatedbyseveralmethodsinandnearKibiareorderedapproximate−
1yas払ilows：
Visitsto apiaries：lT7a17dari／7ia＞simillima＞cn7bro＞anaILf＞
Visitstotreesapsources：mα円（ねデざれね＞cr油ro＞針矧鮎＞加西雄＞扇如拙瑚
Makoto M八丁SUURA
116
Visits to houses：trOPica＞crobro＞ mandorinia＞simE／Iimo＞ maT7dorir7ia
Generalextranidalactivities：mOndarinia＞［rOPica＞ OnGIis＞ crabro＞simi［／
Discover〉・Ofhibernatingqueens：OnaIis＞ trOPica＞silTli／／ima＞ mandarinio＞ Crqbro
Nestdiscovery：∫f研′／′／椚α＞即紺旭＞mめⅦ＞〃矧痛釧偏か＞打叩血
Themarkeddiscrepanciesamongthesedifftrentcrudeestimatesshowhowdifficultitistoaccuratelyestimate
therelativeabundanceofspeciesinagroupsuchas掩spa；Withawidenightradiusandrelativelylowdensity．
Itispossiblethatwemayestimatethenumberofalmosta1lofthecompletednestsof V：sirnillimain
adefinedareabysearchinga11theirmostlikelynestingplacesattheendoftheseason．Thble43showssuch
’払ble43，FltlCtuationsinthenumberofnestsof物asimi！limainfburadjacentareascoverlng300ha．near
Kibi−Cho．1964−1975．
Year
Sub−areaS
A．Mut5ugaWain
Area（ha）
50
Z O
D．協madain
YuasかCho
95
0
1
5
3 10
3
2
0
2
2
4
1
2
3
1 1
2
2
1
1 16
2 17
O
0
00
﹁︼
4
︻ノ
る
0
3
5
9
へJ
﹁▲
5−
受V
0
‘U
八Xu−
4
7・
00
つ、−
Qノ
4
′lU
︵U
′0
く■
1︸
・・・▲
旬U
■・l
′hV
▲ヱ
4
2
つー
︵る
4
3
7
3
00
っ一
﹁▲
A．
′hV
−一
‘1−
′hV
′hU
5
′0
8
4
2
つ︼
＜U
0
﹁︼
7
︷ノ
5
OU
．5．46 誹
へ‘ 鎗V 5
0
l
7
︹J
00
3
′︵V
OO nフ
‘U
・■1
nフ
4
0
0
09
八U
カ
0
00
つJ
2
0
2
2
0
つJ
︵〃
〇
3
Qノ
︻J Q′
4
︵X︶
くノ
5
6
−・一
＊Alllhenestsweretakenartercessationorne5tingactivities．
3
0
0
2
2
4
0
史V
￠
‘V
0 50 4
つJ
8
へJ
5
0
3
9
3
73
Aver色名e 郎】3
23
4 10
0
6
Minimum 2，3る】
Maximum10，058
2
胡崩 50
7
4
0凸ノ
1 1
0
1 1 19
2
0
3
Ⅸ
0．02
6
ユ
300
9
3
PerCO王0Ily＊
0
1 1
0 2
つJ
No．orぐells
1
0
No．nests／ha．
4
之
0
9
Tbtalno．nests
Ob＄erVed
2
】
1
7
C−Yb5himiiれ
Kibトcho
2
3
5
1
i Z
‘V
2
2
4
SO
3
4
B．1bshikalVain
Kanayかeho
3
5
王
八肌
1
3
75
Kanayかeho
7ぬ‡
，糾・65・66，6？・68，69・70 り1りZ り3 ワ4 り5
arecordofthedensityof VsimillimanestssurveyedinfouradjacentareasnearKibi，Wherethenumber
Ofthenestsofthisspeciesisfairlyhigh．Thenumberofnestsperyearvariedonly3．33timesfromtheminimum
tothernaximum，duringthe12years，Suchasmallrangeofnuctuationsofnestsmeansthatthisspecies
isstronglycontrolledbynaturalstabilizingmechanismsregulatingthepopulationsatlowlevels，despitethe
factthatthesecoloniesareestimatedtohaveproducedatotalofl，800−6，000newqueenseveryyear．
7．Discussion
l）Survivorshipcurves
Inhiseornparativereviewonsurvivorshipcurvesforvariousinsects，IT8（1959）suggestedthatbroodmor−
talitydecreasesasparentalcareincreasesandthesurvivorshipcurvestendtobeconvex．Verylowrnortali−
tyatthebroodstagehassofarbeenconfirmedbyseveralworkersinafewsocialwaspsandbees．SpRADBERY
（1973）sho＼＼′edanestimatedbroodmortalit）・Of3．5qDinworkerso＝句叩／agermaI）icE7．1nthehoneybee，
，Jpisme／／昨m，＼Vhichhasachievedhighlydevelopedsocialcarethesurvi＼′OrShipcur＼・eisofthecon、′eX
type，Whichisverysimilartothatofman，andthemortalityofbroodstagesis14．9甲bunderfavourable
conditions（FuKUDÅandSAKÅGÅM王1968，SÅKAGAMlandFuKUDÅ1968）．Theypointedoutthatthecondト
tionsnecessaryforbroodsurvivalinnormalcoloniesofthehoneybeearekeptwithintheoptimairange
by【headult workers．resultingin averylow mortality．
Theresultsobtainedfromthefive掩軍aSPeCiesobviouslyconfirmsthis．However；thevery10WmOト
talityrateinbrooddevelopmentalstagesofworkersisachievedonlyunderfavourableconditions．Insuch
ComparativebiologyortheriveJapanesespeciesor【hegenu5物α（Hymenoptera，鳩spidae）
117
COioniesmortalityfactorsoperatingonthebroodareveryfewexceptfbrcannibalismbytheadultworkers，
beeausetherearefbwnaturalenemiesabletoattackthemindividuallyintheirnest．Ontheotherhand，
nursit唱adultsexualsduringthcreproductiヽ・ePeriodevokesacatastrophicbroodmortalit〉1eVeninfavourable
COIonies，WhichisprobablycausedbyadecreaseinattentionglVentOthebrood．
Thereareessentia11ytwomortality払ctorsoperatingatthisperiod，i息，Cannibalismbyadulthornets
andstarvalion．TheformerisliterallyapartofthecoIony‘fbedback’systemasthemeanstoretaimng
proteinroodsupplywhenfoTaging、、∫Orkerschangetheirmainfoodcollectionfromproteinforthelarvae
tocarbohydrateforadultsexuals．Inthiswayworkersutilizethebroodastheirproteinfbod．Withreduc−
edfbeding，mOStOfthelarvaeremaininthelastinstar；thoughtheycontinuetosupplygrandularsecre−
tionwhichisfrequentlytakenbyaduIthornets，andsuchcircumstance＄Obviouslyaccelerate＄tarVation
Ofthebrood．Finally，thebroodduringthelaterstageofthecoIonycyclearedestinedtobeabortive
OneSintemperateregions，aSalreadypointedin勒ぬteswasps（MATSUUR＾1977）．
Comparedwiththelowmortalityofworkerbroodinnormalcoloniesthemortalityrateoftheadtllt
Stageiscompensatinglyhighwiththecommencementoftheirforagingactivitie＄．IsHÅYetal．（1967）noticed
thelowerlifbspaninworkersof物aorienta蝕havingaccesstothefieldandreportedthatduringthe
first14daysofadultlife，theworkerssufferedamorlalityof8．8甲oinconfined tothenestand42．5甲o
ifallowedtofbrageoutside．lnhoneybees，Ap由me〟腑TqWhichdonotnormallybeginfbraginguntil
thetenthdayorlateちmOrtahtyisalsoveryhighwhentheybeginleavingthehiveonfbragingtrips（SEXIGUCHl
andSAKÅGÅML1966，SAXAGAMlandFuxuDA1968），AccordingtoFuxul）ÅandSEXrGUC川（1966），mOrtality
fhctorsofhoneybeeworkersduringtheforaglngPeriodaredividedintothreecomponents，eCOIogical
haヱards，physiologicalconsumptionduetointenseforagingandtha【duetointensebroodrearingduring
theearlieradultstages，thoughitisimpossibletoseperatethem．
ln掩軍喝WOrkersstarttoieavethenestonfbraglrlgtripstwoorthreedaysaftertheiremergence，and
COntinuetheirfbraglngaCtivitiesusuallyuntiltheirlastdayalive，thoughitisreportedthatnoworkers
Of1lorie（T（a］isleavethenestafteranageof35−38da）rS（1s11＾YetaJ．1967）．Mostoftheforagingworkers
areconsideredtodiefromecoIogicalhazards，SuChaspredationoreventualaccidents，andthosewhich
havenotencounteredsuchmisfortuneswoulddieofphysiologicalseneseence，thoughitisverydifficult
to measurequantitivelytheeffect or5uCh factorso11themortalityby direct observation．
1tseemslikelythatthelon邑eVit〉rOfLifespanin掩岬aWOrkersiscloselyrelatedtothesizeofpopulation
insuchawaythattheIongerthelifespanofadultsthesmalleristhecolonysizeattained▲Itshould
benoted【hatthelifespanoftheworkersof t・1（ropEca，≠′hichhasthesmallestcolonysizeandtheshortest
activityperiodintheJapanese Vaspqisremarkablylongascomparedtothatoftheotherfourspecies．
Fina11yitmustbeemphasizedthatin fespaspeciesthemortalityfactorsaffectingeachindividuaまin
acoIonydifftressentiallyfromthosewhichoperateatthecolonylevel：thelatteroftencausecatastrophic
damageto allthecolony．
2）Seasonalchangesin population structure
TheseasonalchangesinthepopulationstfuCtureOfvespinecoIonieshavebeendiscussedbymanyworkers
（JANET1895，BoDENHEiMERI937，fsHAY1965，IsHÅYetal．1965，MÅCDoN＾LDetal．1974，RousHandAKRE
1978，ARCHLR1982）．L6vGREN（1958）consLruCtedamathematicalmodelmeasuringtheseasonalgrowth
andproducti＼′ityof＼、′aSPCOlonies，uSingparaIneterSbasedonlセspILIaspecies，takenfroIllRJC＝∧RDSand
RICI抽RDS（1951）．SpRADBERY（1971）exemplifiedL6vgren，smodelin Vaspuねspeciesandprovidedmuch
betterestimatesoftheparametersforl／bpzJ／acolonies．AR川ETt（197ユ．1981）alsogaveanaccountofthe
SeaSOnaldevelopmentof TespuねcoloniesinrelationtocharacteristicsofcoIonymembers．
Thcgeneralpatternof、seasonalcolonydeヽ・elopmentin沌叩aischalaCterizedbyslowde＼でlopmentin
theearlystage（MATSUUR＾1973），fbllowedbyexplosivedeveIopmentinthelaterstageandthentherapid
andterminaldeclineaftersexualshavebeenproduced．Thisissimilartothatofmanyspeciesoftemperate
polistineandvespinewasps（JANET18タ5，1903，SpIET王i1948，YosHIXAWA1962）．Concerningthegeneral
patternofthecoIonydevelopment，nOmarkedvariationexistsamongtheJapanese物aspecies，but
SOmeSpeCificdifferencearerecognized（Fig．65）．Thatprincipaldiffbrencesareasfollows：（l）duration
118
MakotoMATSUU見入
ofcoIonycycles，（2）attainablesiヱeOfthematurecolony，（3）timeoftheemergenceofsexuals・Comparing
tbedurationorcolonyc匹1es，tbeco10nialu托orIて′叩∼甜isabout3・5to4months，Whichistheshortest
amongtheJapanese物aspecies，Whilethatof Vsim〟！imaisnearlytwiceaslongat7to7・5months・
Theotherthreespeciesoccupyplacesbetweenthetwoextrernesinthefb1lowingorder：Simj〟ima−mandbrir7jd岬
a77aHS−CrUbTV−1rqpicaAsoverwinteredqueensofeveryspecies，eXCeptfbr V：trppicqbegintobuildtheir
nestsataboutthesarnetimeduringrnid−tOiateMay，thediffbrencescouldresul【fromthefactthatthe
long−CyClespeciesgenerauyneedmoretimeinorderfbrthecoIonytomature，andattainabiecolonysize
ispartlybutnecessarilylinkedwiththedurationofcolonycyclesinsuchawaythatthelargerthetypic
colonysizethelongerthecoIonycycleisprolonged．Asimuartrendisreportedin物uおandDolichov呼u由
species（MACDoNÅLDetal．1974，GREENE1タフ9）and劫触びSpeCies（MATSUURA1975）・Hけ叩加whichis
theveryshorトCyClespecieshasthesmauestcolonysizeamongtheJapanese物aspecies・Ontheother
hand，打ざ∫椚′〃加αWhichislong−1iveddevelopsenomouscolonies，aSlargeasthoseorsome物〟ぬspecies
（MATSUURÅ1980，YÅMANEetal．柑80）．Thecolonysizeが㌣仁和∂和and折椚ク〃血rf乃由ismediumamong
the恥e物αSpeCまes：thenumberofceusinthelargestcolonylneaChspeciesisabout4，500，although
Hぐ招かりCOloniesaregenerauysmauerthan巨椚β〃ぬrg〃ぬcolomies・Thedurationofcolonia用鎚ismuch
shorterfbr VmtbTVthanfor Vmanddrinja：theendoftheco10nyintheformerspeciesoccursinmid−
tolateOctoberwhereasitextendstolateNovernberinthelatterspecies．Thecolonyof VanaHsismuch
smallerthananyother挽甲αSpeCiese㍍eptfbrlて汀叩如buttbeincreaseinnumberoぎcellsisverysmau
duringtheactiveperiod，thoughthisspeciesisratherlong−1ived・
Sexualproductionalwaysoccursattheendofcolonydevelopment，SOthetirneofemergenceofsexual
iscIoselyrelatedtothetimeofcolonyterminationineachspeciesINewqueensemergefromearlySeptem
tolateNovemberinthefbllowlngOrder：′′印加−C和わ代トα〃α払∫油′／肋〝一椚α〃血r加血Justpr10rtOthepro−
ductionofnewqueens，thelarva：WOrkerratioisconsiderablyreducedanddropstoitslowestlevel，below
about2，Oineach挽甲aSpeeies・Suchadecreaseinthelarva：WOrkerratioiscommonlyrecognizedinsocial
waspsandbeesasoneofthemostsigni蔦cantcyclicchangesatqueenrearingtime（R［CH＾RDS1946，SpRADBERY
1965，197l，ARCHER1972）．Thedecreaseinthelarva：WOrkerratioindicatesanapproachtoasuitablecondト
tionfbrqueenproduction．PoTTER（1965）suggestedin掩軍uktthatamajorcontributiontothedecrease
inthelarva：WOrkerratioresultedfromthernarkedincreaseinworkerlongevityatthelaterstageofnesting・
Ontheotherhand，theminimumnumberofworkersinamaturecolonywhichisregardedasnecessary
tosucceedinprodueingnewqueensvariesconsiderablyamongthespecies・Fbrexample・in Vtrppicanew
queenscanbeproducedinaco10nywithmorethanca・20workers，Whileinmostcolonieso
Fig．糾．Schematicrepresentationortherearingtimeornewqueensinthe恥e物αSpeCies・
Col】1Parati＼■e biolog）IOrthc n＼・eJapancscspecies orthcgenuslセ岬a（Hymcnoptcra，Vespidac） 119
thenumberofworkerspre＄entinareproductiveIysuccessfulcolonyneedstobeatleastca．3OOworkers．
Furthermore，itisclearthatthelargertheeolonysize，themoresexualsareproduced，thoughcorrelation
betweencolonysizeandthenumberofnewqueensproducedinthecongenericspeciesisnotmarkedly
Obviousexcept fbr H∫／用f／／f椚α．
Recently，ARCHER（1980）pointedouttheexistenceofanalternativestrategyincolonydevelopmentin
Britishsociaiwasps：‘long−CyCle’astypifiedbyI毎uktv鴫arLsand‘short−CyCle，，e．g．Do／ichove5Pu血
相Ivestr玖anddiscussedthecharacteristicsandtheevolutionaryslgnificanceofthetwostrategiesbyus−
ingacornputermodel．InthecaseoftheJapanese挽甲aSPeCiesitisrecognizedthatthereiseverygrada−
tionofcolonycyclefrom‘shorトCyClespecies’withasma11colony，e．g．V（J叩icq‘long−CyClespecies，with
alargecolonysIZe，e．g．Vsimil／ima．Theotherthreespecies，Vc／Ⅵb／q VanalLfand Vmanddriniahave
anintermediate−CyCle，butoccupydifferentpositionsbetweenthetwoextremes．Consequentlyitseems
likelythattherearingtimeofnewqueenlarvaeissomewhatdifferentineachspecies（Fig．84）aspointed
OutinEuropeanwaspsbyARCHER（1980）．Asmentionedinchap．ⅠII，thefive物aspecieshavesomewhat
diFferentrequirementsfbrsourcesoffood，eSpeCiall〉・PrOtein．Therefore，thedifferenceinthelengthof
thecolonymaybereflectedpartlyinthedifferenceoffbodsources，andthefive物aspeciesprobabiy
reducecompetitiveoverlapwitheachotherforfoodsources．Thisresultsintheiroccupyingdifferentniches．
3）Mortality factorsat colonyle＼′el
IthasbeenemphasizedthatmostmortalitylntemperateSOCiatwaspsoccursinthesolitaryperiodas
reportedin馳ぬtesspp．（YosH王l（AWA1954，YAM＾NEandK＾W＾MACm1975，MATSt）URA1977，MIYANO1980），
in掩spuhtspp．and Dolichovequぬspp．（DuNCAN1939，ScoTT1944，BRl＾NandBRIAN1948，1952）．In
thecoloniesofPo独tes；COIonyfailurerateinthesolitaryperiodisvariablefrom42．6勒（MzYANO1980）
to84．2％（YosHTXÅWÅ1954）bydifferentlocalenvironments．YosHIKÅWA（1954）infbrredthatthesefailures
WeremOStlyduetolossofthefoundress．Ontheotherhand，BRIANandBRIÅN（1952）andARCHER（1980）
noticedthatinDolichov叩uね5y［vestriscolonies，mOStSeemtOfailonaccountofsomeapparentweakness
Ofthequeen．
Asto掩軍ahornetsthedataontheunsuccessfuinestsobserYedinthefieldshowedthatthecolony
failurerateinthethreespecies，Vana晦Vcrvbroand VtfVPi叫duringthesolitaryperiodwasremarkably
highincomparisonwitllthatoftheother［WOSPeCies．V ma17dariniaandlてsimi［／ima．Aspointedout
PreViously，however；thechancesoffindingtheincipientnestsof VtrqpICaWereVerySmallbecauseof
itscoverednidificationinadditiontotheprlmarylowdensity．Thus，itseemslikelythatthereexistmany
morenestsofthisspecieswhichhavediedoutattheincipientstagewithoutdiscovery．Itisprobablethat
Simialrlyheavyearlymortalityduringthesolitaryperiodoccursamongallthesefivespecies．Datafbr
nestsurvivalindicatethehighfrequencyofnestdemiseoccursduringthecooperativeperiodaswellas
atthesolitaryperiod，andthemainfactorforcoIonyfailurecouldbeattributabletothedisappearance
Ofthequeenduringherforaglngtripsthoughitisimpossibletoestimatethecauseofthisdisapperance
indetail．Oncereachingthelaterpartofthecooperativeperiodinwhichthequeendoesnotforageout−
Sidethenest，herrnortalityprobablydecreasesbecauseofliberationfromforaglnltaSkswithhighrisk．
Afterthepolyethicperiod，COlonymortalitydecreasesmarkedlyasLthecolonybecomesestablishedand
StrOngagalnStinvaders．Nevertheless，ahighrateofcoIonydestructionoccurredinthematurenestsof
bothlてsimillimaand Vmandariniaincomparisonwiththoseofother掩spaspeciesinspiteoftheir
formidableabilitytoattackandtoconquerotheranimals，Itisremarkablethatthemostmortalityoccurs
byattackof L／：mandariniainthecaseof11∫i川i／／（maalldbydestructionbymeninthecaseof tて］T柑J卜
dariniq andtheseplayanimportantpartasreguiatingfactors．
itshouldbepointedoutthatIセ軍aCOIoniesseemstoremainatafhirlylowleveiofabundanceinat
leastsouthwesternJapan，inspiteoftheirhighproductivity．lnthecaseof Vsimi／limawhichcanpro−
ducemorethanl，00Oqueenspercolony，ahighrateofreproductionwouldrequlreamOrtalityofmore
than99．9勒tomaintainthesamenumbersofcolonieswhichreachthereproductiveperiod．lnotherwords，
atthemostonlyafbwqueenswouldbeexpectedtosurvivetomaturityinastablepopulationofcolonies
inanarea．ThesametendencylSrepOrtedin掩岬uねwasps，tOO，anditissuggestedthatmostmortality
Makoto MATSUURÅ
】20
OCCurSbefbrenestsarebuiltorduringtheperiodofcolonyinitiation（BRIAN1965，SpR＾DBERY1973，
ARCHER1980）・Therefbre，itisassumedthattheremustexistaheavymortalityof Tkspaqueenswhich
mayoccurduringtheperiodfromthedispersalofnewqtleenSfromtheparentnesttothefoundingof
new coloniesin sprlng・Possible factors affecting smallvespine populations such as始軍uねand
Dolichov叩uhthavebeendiscussed，eSPeCiauyclimaticfactors（LATTER1985，BEIRNE1944，FoxW）l．SON
1946，D6HRING196O）・Theseobservationsindicatedthatthemeteorologicalconditionsdidnotplayadomi−
nantroleinqueencontrol，buttheya11consideredthatsprlngWeatherwasimportantasafactorinqueen
COntrOl．
Factorssuchasnaturaienemies，physioIogicaldeath，etC・mayCauSealossofhibernatingqueensto
acertainextent・However，thecauseofdeathofthebulkofqueensofvesplneSPeCiesremainsuncertain
aswasdiscussedfbr物uiaspp・（SpR＾DBERY1973，ARCHER1980），thoughintra−andinterspecificcom−
petitionamongqueensfbrnestsitesinsprlnglSlikelytobeoneofthemostsignificantfactorsinpopula＋
tionre如ation，eSpeCia11yduri喝yearSOrabundantqueens，leadingtou5urpation（N】XON1936，BR】刷
andBRlÅN1952，MÅTSUURÅ1970，SpRAD8ERY1973）・ARCflER（1980）suggestedtheexistenceofselトregulating
mechanismsin挽軍〟ぬv塘α血whichcausedatwo−yearCyCleorwaspabundanceandscarcityinEn釦and．
Hepointedoutthatalinkbetweenqueenqualityandcompetitionfbrnestsitesinsprlngplaysthemost
importantroleasalikelyselトregulatingfhctorlThenumberofmaturecoIoniesin掩軍aSpeCieswasrelatively
COnStantirrespectiveoftheirhighprodtlCtionofnewqueensascomparedwithviolentfl11Ctuationinnumbers
Of掩甲u］acoIonies，Whichshowa10−42−folddifferencebetweentheextremesofscarceandabundant
years（NICHOLSON1921，BEIRNE1944，FoxWILSON1946，EDW＾RDS1974，1979，ÅRCHER1980）．
lnthissectionIhavediscussedmainlythelossofcoIoniesin掩軍aandrefbrredonlytothosecolonies
Whichhavepassedtheirvulnerableincipientsoiitaryperiod・Hovnvenitisprobablethatthereexiststabilizing
mechanismsfbrtheentirepopulationof掩軍aSpeCies・職understandthisitisofgreatimportancethat
anaccurateestimateofthenewqueensinthesixrmonthsnon−neStingperiod，thoughthereislittleimfoト
mationonthissubject．
V］．GENER＾L D［SCUSSlON
ThesubfamilyVtspinae，CertainlyoccupylngaSummitofvespidevolution，COntainsonlyfburgenera：Prv−
V叩払か0／ねカove軍〟毎物〟ねα〃d掩軍α，
Thefirstgenusisconsideredtobethemostprlmitive，andthethreeknownspeciesoccuronlyinSoutheast
Asia（VECHT1936，1957）・Themostsignificantbehaviouraleharactersofthisgenussummarizedinmyrecent
WOrkinSumatra（MATSUURA，unPub・）are：（1）typica11ynocturnalactivity，（2）nestfbundationbyswarming
Withasinglequeenandseveraldozensofworkers，（3）verylownumbersofqueensproduced（＜20queens
percoIony），（4）lowvaiuesfbrthelarva：WOrkerratioatanypointinthecoIonycycle（＜3．0），（5）heterogeneous
queenproductiononasinglecomb，（6）specialqueen−rearingce11swith耽tremelyprolo喝edcellwa恥（7）
treebranchnidification，（8）thetoughandcomparativelypliablenestwithacompleteenvelopeconsisting
Ofafewfblt−1ikelayersonthetopsurfaceandnumeroussmallscallopedstructuresintheupperhaif，and
SeVerallayersofsmooth，broadoverlappingcartononthelowerhalfinamatrixofdense，rigidcarton，（9）
鮎blece11wallscrapingbymaturelarvae，（10）presenceoradistinctroyalcourtorworkers，SurrOunding
thequeen，（11）irregulardistributionofbroodonacombcausedbyutiliヱationofthesamecellseveraltimes，
（12）multipleutilizationinthesamequeencells，（13）strictlypredacious，Carrionnotutilized，（14）longcoト
Onyduration（7∼10months），butnotperennial，（15）ratherlargecoIonysize（Ca，1，500workersandca．
5，000ce11satpeakinきケove軍α刀0α〟mα），（16）presenceoぎnyingabilityinthequeen血ou由outherli托time．
Someofthesecharacters，eSpeCial1y2，8and14canberegardedasadaptationstoenvironmentalfactor
OftroplCalreg10nS‥SeaSOnalityandnaturalenemies・TheswarmlngmOdeofcolonyfoundationisalmost
exclusivelytropical，andsomepeculiaritiesofnestingbiologyenumeratedabove，eSpeCially3∼5，15and
16，appeartObeconnectedwiththemodeofcolonyfounding．Thishabitisinterpretedasanadaptation
toahighlevelofantpredation（JEÅNNE1975）・Ontheotherhand，thedescriptionsgivenabove，eSpeCially
ComparativebioIogyofthefiveJapanesespeciesofthegemlS掩3Pa（Hymenoptera，Vespidae）
121
1and8，SuggeStSOmebionomiccharacters，SpeCifictothisgenus，Whichareofsomeinterestforthenoctur−
nalactivities，1ikeApoicainSouthAmerica（RICHARDSandRICHARDS1951）．Thesebehaviouralcharacters
asindicatorsofvesplnephylogenyshowthatthis5mallgrouplSadaptedfbrlivir唱intropICalreglOnSaS
the most specializedgroupamongtheVespinae．
Both物u／aandDoHchov叩uねaredistributedinthenorthernhernispherewithacompleteadaptation
to temperate reg10nS and are not foundin troplCalregions．Comparing the two genera，物uIaand
Dolichov叩u毎itis8SSertedthatthefbrmerasagroupISrnOreadvancedthanthelatterinsomecharacteristics，
thoughthecomplexismonophyleticwiththeancestorsofthegeneraprobablyevoIvlngfrom掩軍aStOCk
（＼／EぐHT1957，RICHARDS1971，YAM＾NE1976）．Somespecificandgenericdiff七rencesbetweenthemarepointed
OutinThble44，basedupondatafromtheJapanesespecies，thoughcomparativebiologyofbothgenera
Thble44．Speciesgroupsandsomebiologicalcharacteristicsofthetwogenera，柁spu／aand Do／ichol
inJapan（M＾TSUUk＾1981and unpub．data）．
挽甲〟ね
β0／如○りだ叩】J／打
肌理稽血血如乱Id∫β加〃血
J押gdぬ
nc peak
Approximale
C（”onydumtion
5（〉0−l．500
3．5・4．0
川なαrお
usu＆】lysubterranean．somelimes
、車・・・ヾ：−：、−∴．−ニ∴ニ；、ミ；：∴．い、‥ ∴
No．orceまIs
dJJ血メ
ガ〟V′ぐ叩∫
PrOl亡亡l亡d or a亡rial
5（）0・l，200
4．0・5．5
3．αX）・17，∝旧
8．0・9．0
5，（X氾・け，（郁
usua11ysじblerranぐan．
iO111⊂til11亡S P101亡亡t亡d
3，000・10．∝沿
8．0・9．0
∬力作れぐ紺
る．8・7．5
308・2．500
4．0・5．5
（mor再1S）
DeぐIirle
N0．OrneWqueenS
PrOducedper
COloIly
m払A喝．tOmld・Sep．
30・150
earまyDec，
1ale艶p．to mid・Sep暮tO
earlyl）ec．
1aleOcし
earlyS叩・tO
mid・S叩．
封ト・ほ0
ヰ00・1，500
500・2，5（旧
300・l．M
50 -4000
nlid・Aug・tO lale決p．10
mid・Oct．
queen王IeSt
Pedic亡箋
A di5ぐabove
lhe comb■
Envelope
Malu托neSt
Su5PellSOria
Envelope
Preydiverぷily
lwis‡edwllhaha門島lr唱 IWisledwllh乙I tWisledwit‡1a 抽王8tモdwi抽a lwisledwitha
5heet
h8ng】n名言hごet hamg一喝Sheet lla】1glngSheel hangtngsheet
absenl
absent
present
present
Itrfscnl
lwistedwitha
han夢In8Shee【
presenl
‡級minaralldpliab蔓ぐ 1鋸mi！lar8肌dpliめ1elaminar∽d fねgilelanlinarandぬ紘i‡e】amiれ甜卸d触gile lamin訂鋸Idpli曲Ie
butlress‡ike
bultresslike
如IinaraれdpliめIe Iaminarandpliable
Stricllypreda10r＄
cordlike
cord独e
5Caliopedand sぐalIopedand
汁agile
rr乱写ile
cord‡i毎
5ぐa】王opeda11d
bothlypespresent
1an】i11ara11dp‡i油1e
rraさi‡e
油Iypredators Pr慧：；ご…uS Pr慧：…ご：uS Pr慧…ご…u5
strictly predators
†ar【er Y入MA郁andM八紘INO（1981）
hasnotyetbeenstudiedindetail．1ngeneral，membersof物uぬexceptfbr掩軍u由schrenckiihaveiarger
COIonies with alonger co10ny duration，and have developed an omnivorous habit，Whereas members of
Dolichov呼uねand fespu血schFmCkiihavesmallcolonieswithashortcolonyduration，andaTeuSuallycon−
Sideredtobestrictlypredators．Itseemslikeiythatinthesegroups，thecoIonysizeattainedatpeakislinked
bothwithcolonylifbspanandpreydiversity．
Similargenericcharacteristicsincludingthoseofthe掩軍uhlniPgrouparereportedfromtheNorthAmerican
SPeCies（DuNCAN1939，RousHandAxRE1978，GREENE1979，ARREetal．，1981）andtheEuropeanspeciesO（EMPER
Makoto MATSUURA
王22
andD8HRINGl％2，SpRADBE鼠Yl卯3，EowÅRDSl粥0，ARC＝ER1980，1981）．ARCHER（1980，1981）stressedthat
thestrategiesbetweenthemreflectbasicintrinsicdifferencesmainlyofagenericori郎n，andthedevelopment
Oflongandshortcyclespecieswouldreduceeompetitionbyailowlngthepeakcolonialpopulationsoflon
andshortcyclespeeiestoappearatdifferenttimesoftheyearsothatpossiblecompetitionbetweenthem，
e・g・fbrfbodresources，WOuldbereducedalthougheachspecies，inturn，WOuldexploitthatresource．Possibly
hisinterpretationisusefulinexplainingtheevolutionofcoIonycharacteristicsbetweenthetwogene
referenceisnotmadeconcemingthepreydiversitybetweenthem・Ofparticularinterestisthatinbothgenera
SOmeSpeCiesleadapermanentsocialparasitic肋．InthecaseoftheJapanesespecies，I卸u血austriacaand
加／油ov叩〟ね〝血蛮血αdonotpossessaworkercastebutaredependentuponhostworkersoぎ打∬カ柁〃ぐただ
and D norvegicoidesrespectively，tOreartheirbroodoFsexuals．
TheIastgenus，Ik5Pq・hasseveralspeciesintroplCalreg10nSbutthegreatestnumberofspecies，aSWellas
thegreatestdiversitylnCharactersarefbundintheeasternHimalayasandsouthernChinaandtheseareasare
regardedastheoriginalcentreofthisgenus（VECHT1957）．Someofthespeciesarefbundtobedominantin
temperater喝10nS，decreaslnglnnumberorspeciestowardsthenorthernandwestern王imitsordistribution，
incontrasttothatofl吻uぬandDolichov呼uk7ineasternAsia（Y＾MANEetal．198O）．InJapanboththe
genera挽甲〟ぬand伽／励ov叩〟ぬaremoreborealorlivemainlyinhighaltitudes，ingeneraldistribution，
Whereas噸aarerareorabsentinhighaltitudes，andthefivecloselyrelatedspecies，Whicharetreatedin
thepresentpaper，COmmOnlycoexistinlowmountainsandfbrestsandoccupythetopranksamonglnSeCtS
andothersmallanimalsin sou【hwestJapan．
1tisofparticularinteresthowthesefivespecieshaveattainedastat）leco−eXistence．Themostlikelymodifica∼
tionsoradaptationsinnicheexploitationareasfbllows・First，thesefivespeciesdi胎rfromeachotherinnesting
Siteastheyareclassifiedintotheopenplaceandcoveredplacetypes・Vanaband VsimjHimabelongto
thefbrmertypewhileVsimi〟imachoosesomesites，typicalofthelattertype・VtrqpLCaandVcrubTVPrefbr
tonestonlylnreStrictedcavitiesorinamplespacesbothaboveandunderground，Whereas fてmandark7iais
exclusivelysubterranean・Thesediffbrencesareverysubtletoavoidacompetitivedisplacementandhaveevoiv−
edavarietyofspeciauzationsinnestarchitectureandmodificationsofthenestsite，SuChasrelocationofnests
Performed b〉・both Vsimi／／imaandIてm7bro．
Secondarily，thefiveI句paspecieshaveachievedaco・eXSistenceintheirftedingrelationshipsbysignificant
difftrencesarnongtheminsuchawayastobeeitherhighlyspecializedorverygeneralizedintheirfbod
Pre飴rences・VtrwicahasevoIvedtospecializeonlyonthebroodofthepo最stinewasps，Whicharehighquality
fbodsources・However；thisstrategylSpreSumablyonlyprofitabletothespeciesofthesmaupopulationtype
SuChasVtrqp娩notrequlrlngalargequantityofanirnaldiet．Ontheotherhand，Vc7VbFDisasemi−SpeCialist
Whichishigh1yspecializedt）ehaviourauyfbrutilizlnganyCicada，Whilekeepinganabilitytohuntawidevarie−
tyofinsectswhenthepre知redfbodsourceisrestrictedindensity．Both Vanalisand VsimiHimaarenon−
SpeCialistsutilizingmanykindsofliveinsectsandspiders，thoughthereseemstoexistsomespecificpreたrences．
Inthesetwospeciescompetitionfbrresourcesisreducedatleastinpartbythefactthat Vana胎breedsexuals
eariierthan Vsimi肋n偽inadditiontothedistributionofscatteredfbodsourcesandtheirlongfbraglngdistance
（1∼4km）nightradiusinbothspecies（MATSUURAunpub．）．Vmandariniq，Whilekeepingitspolyphagoushabit，
evoIvedaspecializedwaytoattackinmassthecoloniesofsocialbeesandotherwasps，includingthoseof
COn＄enericspecies．ThisresultsinadisplacementoftheecoIogicalnichesbetween Vmanddtiniaandother
CO喝enericspeciesissuchawaythat打椚α〃血r∼〃ぬa晩ctsthemintwoways；aSaCOmpetitorinseekingco汀ト
monfbodsourcesandasapredatoちbeingthemostimportantnaturalenemyatthecobnylevei．H椚釘ト
dariniaowesitssuccessinthestrategynotonlytoitslargebodyandcolonysIZeS，butalsotoefficientgroup
Predation，WhichistheonlycaseofgroupfbraglnglnSOCialwaspsthusfarknown．Thus，thesedifferences
infbodpreferenceamongthefivespeciesresultinreducedcompetitionfbrfbodand主nutilizingthesesources
atslightlydifftrenttimesoftheyear，Eachspeciesreachesitsgreatestneedforproteinfbodtocoincidewith
theoccurenceofitspreysothatcompetitioniseffectivelyeliminated・Therefbre，itissuggestedthattherearlng
timeofthesexuals，Whichneedthegreatestquantityoffbodstofbedthem，issegregatednotonlybyseasonal
timebutalsobylengthofcoIonycycleasshownschematica11yinFig．84．
ComparativebiologyofthefiveJapanesespeciesofthegenus掩5Pa（Hymenoptera，Vespidae）
123
Theprincipaldifftrencesare：Vt／叩icaisashort−CyClespecieswiththesmal1estcolonysIZe，Whereas Vsim肋a
isalong−CyClespecies、、・iththelargestco10nySize．Theo【herspecies．レ：crubrql（ano／isand L，：
OCCupyintermediatepositionsbetweenthetwoextremesbothinthelengthofcolonycycleandcolonysize，
Thus，thefiveⅥspaspeciesseemtooccupyecologicalIydifferentnichesinsuchareas（職ble45）asnest
Site，fbedinghabitsandcolonycycle，WhichtheyhavecoexistedthusfhrpermitcoexistenceinsouthwesternJapan．
Thb】e45・tnterSPCCirjcrankingamongtheriveJapaneseIセ5PaSPCeies．
Species
∫i刑∼／／∼∽α α〃クJね
ぐJ甘み／・0 椚α〃dα′∼〃ぬ
Number of nests studied
FirstappearanceorqueerlSinsprlng
Firsteme曙eneeOぎworkers
Nes（leavingbynewqueens
First deathoffoundingquccns
Disintegration of colonies
Size orcel韮s
Maximunlnumber ofcellsin nest
Numberofsexuals percoIon），
Ag托SSivenesstowadsman
Interspecific dominance
Abundance rneasured by：
a）visitstoapiaries
b）vi5itstotreesapsotlrCeS
C）visitstohouses
d）general飲tranidalactivities
e）discoveryorhibernati咽queenS
f）nestdiscovery
SUMMARY
ThepresentstudyaimstoclarifythespecificdiffererlCeinbionomiccharactersofthefiveJapanese掩spa
SpeCies．Vsimi／Iima，Wn（hop（emC刷ERON，Iてana［LfinszLlarねD＾Lし＼ToRRE，IてcrabroJ7avq舟Seia（aC＾MERON，
IJ：mandariniajqponicaRADOSZXO、＼・SKIandIく（rOPicGPu／chraBul・SSON，＼Vhichcoexistinlowmountainareas
OfsouthwestJapan．Thelifbhistory，neStinghabits，feedinghabits，divisionoflabourandpopulationdynamics，
aredescribedprlmarilybycomparlngthemwitheachother．The蔦eldworkwasmainlycarriedoutinand
nearKibi−Cho，Whkayama，SOuthernJapanduringtheperiod from1964to1975．Atotalof6ユ2nests＼VaS
Observedinandnearthestudyarea，andconsistedof288nestsof Vsimi！limち1640f Vana晦800f V
Crabrp，560FIイma［Tdari／血and340fレ：（ropica．Results obtainedareas fbllows：
1）Theannuallifecycleofthefive tk5PaSPeCiesconsistsofthefbllowingsixperiods，thoughalittlevariation
indetailsofthecycleoccursamongthem：（1）pre・neSting，（2）solitary，（3）cooperative，（4）polyethic，（5）
reproductiveand（6）hibernatingperiod．Hibernatedqueenscomeouth・Omtheirhibernaculaduringthe
period FromearlyApriltoearlyJuneinthe fb1loIVi11gOrder：SimiI／ima−mandarillia＝m7bTD−aJJaILs−
（TPPica．Allofthespecies fbundtheircolonieswithaslnglequeenwhichconstructsanaverageof35−
40cellsandrearslO−151arvaeuptothepupalstagebyherself．Thefirstworkersemerge魚・OmJuneto
Julyinthefbuowingorderwithadifferenceof5∼6weeksinpeakdatesbetween Vsim肋aand Vtrqpica：
SimiHima−C7ttbTV−ana貼−mandarinia−trqPica．Thequeensare5tiuinvoIvedwithbothextranidaland
intranidalactivitiesfbrawhileafterworkeremergence．Thiscooperativeperiodlastsfbr2∼8weeksand
itslengthisinverselyrelatedtothetypicalcoIonysize．Asthenumberofworkersincreases，thequeensof
a11species，Otherthan Vt7tPicabecomerestrictedtoovlpOSition，leavlnga11theothertaskstotheworkers．
124
Makoto MATSUURÅ
Thisexpansivestagelasteduntilthetimewhensexualsareproducedandthedurationofthereproductive
periodlastsaboutamonthineachspecies．Thedisintegrationofthecolonyordinarilytakesplacefrom
earlySeptembertolateNoヽ′emberinthefollo≠Iingorder：（rqPica−C／Ⅵbro−una／Lf−mandarinia＝S（lT7iI［ima．
Thequeenshibernatesslnglyinhibernatingspaceswithinthesoilorrottenwood．
2）Nestsitepreftrenceisclassifiedintotheopenplaceandconcealedplacetypes．1てana／isandlてsim（／／ima
belongtothefbrmerand V：（rqpicq VcTt7broand Lてmandariniatothelatter．ThequeensofIJ：simi／／ilT7a
and V c7Vb和tendtoinitiatetheirnestsinnarrow，COVeredplacesorundergroundcavities．Whenthe
nestbecometoolargeaftertheemergenceof＼VOrkers，the、Vholecolonymovesane、l・，mOreSPaCioussite．
3）Theshapeofthecompletedqueennestwasdividedintothethreetypes：bowトshaped（V trwicaand
Vmandarinia），balトshaped（y：crvbroand Vsimi［lima）andnask，Shaped（Vana抽）．Thegeneralshape
Ofmaturenestsvariesbetweenthespeciesoftheopenpreftrenceandthoseoftheconcealedplaceprefbrence．
Intheformergroup（V：simi／IilT7aandレ：ana／is）theenve10peisusuallythick，COVeringthecombscom−
pletely．Indevelopednestsofthesespeciestheenvelopesconsistofmultilayeredshe11−1ikesheetsencIoslng
man），airehambersamongthem．Inthelattergroup（t／：cmbltLIてIrPPicaandIてmE7ndari17ia），theenvelope
COnSistsofonlyaftwplate−1ikesheetswithimperfbctairchambers，andthelowestcombisalwaysexposed．
4）Thefoodsof挽岬aCOnSistoftwowe11difinedclasses，Carbohydrateandprotein．Thefbrmerareobtained
fromtreesap，honew・dewofaphidsandpsyllids，flowernectar；muShroomandsyrupjuice．Themost
importantsourcefora11theI包甲aSpeCieswasthetreesapexduing什omaperturesoflivingtrees．Both
intra−andinterspeCificaLlyadistinctlineardominanceorderisrecognizedamongthehornetsvisitingtree
Sap．lnterspecificrelationshipsamongbothqueensand＼＼′Orkersatfoodsourcesareasfollows：mandari（1ia
一打／旧／f∫一亡何か●0−∫〟〃〟〟／ナ和一什0〆ぐα．
5）Preypreftrenceof fespadifferedcharacteristicallywithspecies．Vsimillimaand Vana［ishuntawide
rangeofinsectsandspiders．VcfVbroisregardedasaserni−SpeCialist，prefbrringvariouskindsofcicadas，
andishighlyspecializedbehaviouranyfbrcapturingcicadas．VttwicaisatyplCalspecialist，beingalmost
exclusivelydependentonthebroodofpolistinewaspsfbrtheirproteinfoodandthelifecyclewassyn−
Chronizedwiththatofthepreywasps．Vmandariniaattacksinmassthecoloniesofothersocialwasps
andhoneybees，pOSSeSSlnganabilitytohuntawidevarietyoflargeinsectsandspiders．Grouppredation
COnSistsofthreephases，hunting，Slaughterandnestoccupation．
6）Thequeenof挽甲ahornetsdonotengageincellinitiationafterworkeremergence，thoughshestiupar・
ticipateinintranidalactivitiesforawhile．
7）Any掩岬aWOrkerwouldperfbrmanytaskwithinafewdaysafteremergencewithoutalabourschedule
depending onagedistinctionas kno、＼・nin thehone），bee．
8）InthecaseofVana胎workerovipositedafbwdaysafterthedeathofthequeen．This‘substitutionqueen’
wascharacterizedbythefbuowingpoints；（1）actingastheprimaryorsoleegg−layer，（2）lossofhairs
Onthegenaandmandibularbaseandpolishedheadasinthequeen，（3）cessationofallextranidalac−
tivities，（4）superiordominance，（5）receivingagreatdealoffbodn’Omtheotherworkers，and（6）remarkable
PrOlongation oflifespan．
9）Tbtaldurationofbrooddevelopmentfrorneggtoadultvarisontheaveragefrom30．8daysin7（simi［［ima
to40．1daysin‡くmandarinia．Thereisatendencyforthelargerthebodysizeinadulthornets，thelonger
theduration becameamong the fivespecies．
10）Thesurvivorshipcurveduringthepolyethicperiodisofaconvextypeineachspecies；remarkablylow
mortalityduringtheimmaturestagesandhighmortalityafterthecommencementoffbraglngaCtivities
in the adult stage．
11）Aftertheemergenceofsexuals，highmortalitywasobservedthroughoutaudevelopmentalstages，paト
ticularlyeggand5thinst叫WhichwasprobablycausedbyadecreaseinattentionglVentOthebrood．
12）Thegeneralpatternofseasonalcolonydevelopmentinthefive掩spaspeciesischaracterizedbyslow
developmentintheearlystage，fo11owedbyexplosivedevelopmentinthelaterstageandthentherapid
and terminaldeclineaftersexualshavebeenproduced．Thecoloniallife oflて（l’OPicais about3．5to
4months，WhilethatofVsimi伽ais7to7．5months．Theotherthreespeciesoccupyintermediatepositions
ComparativebiologyorthefiveJapanesespeciesofthegenusレをspa（H）′menOptera，Vespidae）
125
betweenthetwoextremes，inthefbllowingorder：Simitljma−manddrinia−ana［is−CTqbTV≠t噸ica．
13）Comparingthecolonysizeatpeak，thernaximumnumberofcellswasca．300in Vttwicqca．9OO
inlてa／！a／is；Ca4，500in Fイ／71E7ndariniaandIイcmbro andca．10，000inIイsimi［／ima．
14）NewqueensernergedduringtheperiodftomearlySeptembertolateNovemberinthefollowingorder：
tropic（トCmbro−ana／is−Simi／／ima−mandarinia．Justpriortotheproductionofnewqueens，［helarva：
WOrkerratioisreducedtoitslowestlevel，belowabout2．0，ineach掩軍GSpeCies，
15）Theminimum numberof＼l′Orkersinamaturecolony＼Vhichisconsidered as necessar）・tOSuCCeedin
producingnewqueensvarieswiththespecies．In Vt7WtCatheycanbeproducedinacolonywithmore
thanca．20workers，Whilein Vsimi沼macoloniesthenurnberofworkerspresentneedstoreachatleast
ca．300workers，
16）Themaximumnumberofscxualspresentpereolonyvariedfrom56newqueensand27malesinaレ：
（rqPicacolonytol，692newqueensandl，123malesina V：simi［／imacolon）・andisrankedassimi／［i
＞maJ7dari／血＞m7brv＞G，7aIis＞trqPica．IIlgeneralitisrecognizedthatthelargerthecolon）r，the
moresexualsareproduced not onlyin thesamespeciesbutalso amongthe fivespecies．
17）Mortalityfactorsworkingmainlyatthenestlevelaredividedintofivemaincauses，（1）disappearance
Ofqueen，（2）naturalenemies，（3）humaninterference，（4）climaticfactorsand（5）unknowncauses．The
endof t／＆spaco10niesoccursveryfrequentlyduringthesolitaryperiodandfollo≠・ingcooperaliveperiod
Whenthequeencontinuestoforageoutsidethenest，Whichisattributabletothedisappearanceofthe
queen．Theheavypredationby VmandariniGreSultsinthegreatestproportionofalltheunsuccessful
nestsatmaturestageineach掩spospeciesexceptfor VmE717dartnio．1nthecaseoflてmandarinia．destruc−
tion by beekeeperspla）・Sanimportant roleincolony mortalityofmaturenests．
18）Annualfluctuationsinthenurnberof Vsimjllimacoloniessurveyedinfouradjacentareascoveringca．
30Oha．gaveanestdensityofO．Ol∼0．03nestsperha．during12years，andthetotalnumberofnests
Varied汁om3∼10peryear」
19）Amechanismtoexplainthecoexistenceofthefive VhspaspeciesinsouthwesternJapanisdiscussedfrom
theviewpointoftheoccupationofdifferentecoIogicalnichesincludingnestsite，foodpreftrenceand
COlonycycle．
＾CKNOWLEDGEMENTS
IwishtoexpressmysinceregratitudetoProftssorSh6ichFSAK＾GAN10fthelnsitututeofLo＼Vltmperature
Science，HokkaidoUniversityforhiscontinunousguidanceandencourageme？tthroughoutthepresentstudy・
Cordinalthanksare also due to Mr．Robin EDW＾RDS Of Rentokit．，East grinstead，Eng）and，Dr．Michael
ARCHLROfCollegeofRiponandlbrkSt．John，EnglandandMr．RobertJ＾COE）SONOfEastCarolinaUniver−
Sity，U．S．A．，fbrtheirreadingandcorrectionofthismanuscript．IalsothanksMessrs．ShigeakiMrYATA，Noboru
HAY＾SHZ，NoboruYAMAMUR＾andMamoruEG＾WAfortheirkindhelpinthecourseofthepresentstudy．
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摘
要
軋本塵スズメバチ属V勾細（Hymenoptera，Vespidae）5穂の比較生態摩的研究
松
棚
減
水油文でほ，日本の函南暖地の低山地から平地に同所的に生息する5糀のスズメバチ属のハチ執すなわち，キイ
ロスズメ′け陣中沈痛拙那詔涌旬柚■αCAME】モON（以下キイロ），コガタスズメバチV毎細川激減な油川血豆
DAl−l−AToRRl；（以下コガタ），そンスズメパチ怖坤αr相方roββγ石蕗∫r∼〟gdCAMERON（以下モン），メ￣オスズノバ
チV如か＝肌用d打沈ね」可矧扇用RADOSZKOWSKl（以下オオ），ヒメスズノバチ佑車αgr呼fⅢ♪〟たあ和BuYSSON
（以下ヒメ）について，各俺の生活史，営巣習性，採餅習性，分業，個体群の動態などを明らかにし◆これらの穐の典
存の轍構な考燦したものである。
調査ほ1964年より1975年までの12句澗にわたって和歌山県有l卦酢畜㈱町とその周辺の地域で行なわれ総計622娘の
スズメパチ属の毅を観察の対象としている。
1．生活史
生活史の質的変化を域虫の社会ぎ袈署練な中心に区分するとつぎの6段階すなわち，（1）肝営巣胤（2）女‡Eの単独償巣
胤（3〉女王と働きばちの共伺常数j軋（4）分業期，（5〉繁麹カスト疲出j軋（6〉越冬期となる。
越冬後の女ヨ三は4月上旬より6月上旬の問に，キイロー勇一オ＝そンーコカ′ダーヒメの順に出現する。各棟とも朝腹
創設で，受瀾したメス（＝女三E）のみが創巣し，星路地営巣i捌こ35∼40房なつくり，そのうち10∼15頭の幼虫な単独で
育てあげる。臆初の働きばちほ6∼7月に，キイローそンーコカ■ダー【勇一オーヒメの順に羽化する。共同常数き抑ま2∼
8週間で，その長さほ椚山卑射こそれぞれの種のコロニ岬サイズと関連があり，卑小コのような大規模常数彗璧の機では短
かく．コガ■タやヒメのような小規模営巣型の簸では長い。働きばちの個体数の増加にともない，ヒメを除く他の4種
でほ，女ヨ三ほ産卵活動に専念し，分業期の後半にほ，働きばちが女王をとりまくローヤルコートが出現する0オスと
新女王ほコロニーの働きばちの個体数が戯多状態に達した時より約1ケ月以上にわたって産出され交尾ほ狼外で行
なわれる。営巣活動の終息ほ9月上旬より11月下旬までで，ヒメーそシーコガターカ■オーキイPの順となっている。
2．営巣習性
営業場所は開放空間型（コガタ），遮厳空間型（そン，ヒメ，オオ），雨空間型（キイロ）に区別される。キイロと
モンは遊敷当初は狭い遮蔽空間を選好する傾向があるが，働きばちの羽化後の7−8月に，より広い遮蔽空間（そン）
および開放空間陣イロ）へコPニ「が移住したのち新女王の産出を行なうことが多く，営巣末∋臼jの巣でほキイロ更胤6
％，そン53．5タすが引越巣であった。
巣の手鍾造は，女王巣の場合，外被の形状により3弘すなわち，月Il型（ヒメ，オオ），椀型（キン，キイp），徳利
型（コガタ）に区別される。成熟巣では，開放空間選好機のコカー●タとキイPでは，外被は貝殻状をした多数の空気室
祀・鶴城されただ1個の側部の出入l二王を残して敷盤全体な完全に被護するが，避赦空間選好糀のそン．ヒメ，オオで
は，外紗ま不完全な空気室をもった数枚のシートで徴求され，敢下段の巣盤が翻こ露出して出入甘を北ねる。
3．採餅習性
炭水化物源ほ5種ともに共通し，樹液，半麹l芸l昆虫の1〕儲，花嵐熱巣，キノコ（シラタマタケ），空椰や空びん内
に残ざれたジュ｝スなどで，汲も愈斐なものは樹液とキノコである。
蛋！ヨ源となる獲物とその狩猟行動にⅠ摘鋼絢爛澗憐がみられる。キイロとコガタほ「何でも屋」で，前者ほ8日射
穐以上の昆虫やクモな狩り，そのうちの60％は双通日の成虫で占められたが，後者ほ8日53穐以上でそのうち双趨目
成虫37．9％についで膜趨目の有則類成虫も32．2タgを占め，アシナガパチ亜料やスズメパチ亜料の種も含まれる。そン
は放物の95％以上がセミ頬によりて占められるが，大型のトンボ，バッタなども揃える能力なもち「肇技研当家」とみ
なされる。ヒメほ灘劇的な「矧−1う家」で，アシナガバチ亜利・の各種の巣な袈りて，その幼虫と桶の体液のみを摂取す
ComparativebiologyorthefiveJapanesespecie50fthegenu5掩軍α（Hymenoptera，Ⅶ5pidae）
る0オ弓−ほ縫独ではドゥガーネブイブイなどの車型のコガネムシ類なゆ心をこ，多肉で動きの鈍い昆虫やクモを狩る「準
酢豚」であるが，同じ磯の働きばち矧訓カミ，同属他種のスズメパチ，クロスズメパチ胤ミソバチ属などの社会性
ハチの巣な射い・相手を全滅させたのち幼虫や蛸な鯨腐する習性な発適させている。これが，明治以掛こわが閤へ噂
人されたセイヨウミツパチに対する旅団攻撃の磯憫とみなされ単独捕食期，鍛聞殺艶聞き占領鯨登瀾に区別して聯
析した。
4．分業
女ヨ三は働きばちの羽化後ほ，外役，ついで内役の順に引退したのち産卵に尊念する。キンの女王の場合，巣づくり
行動には，働勘ばちの出現複成紋服闇にわたって参加したが，この間に（1）背筋の新設，（2）みずから野外で集めた巣材
による外被や膠璧の仲良，刷外磯暑かじりとって得た巣材による房璧の仲良（4凍材を集めずに，働きばちが伸ばし
た波後の湿った房誓藍の附【しl二（砕く伸ばす）の順に軌とする。
働きばちの鉦齢と仕事に関してほ，5種ともに羽化後2日巨iより各種の内役を行ない，3∼4則］にほ定位飛行の
のち，すく−−に娘軋食物，水などの採集活動に従射し，老齢化しても，巣の内外のあらゆる仕事及こなす能力をもつ。
女王の亡失巣でほ，数日後に働きばちの一部の卵巣が発適し，女王の代位となる胴体が出現する。コガタの場合
「代位女王」は次の特性なもつ。1）唯一一の，またほ憺豪家泣の産卵老としてオスな虔出する，（2〉分業期の女王にみちれる
ような体毛の脱乳および体表面が油じみた光沢なもつ，（3）あらゆる外役への不参れ（4他の働きばちに対する優位
行動，（5）他の働きばちからの霧夏祭な食物摂取，（6）肇命の著しい延長。
5．値…極押の助態
働きばちの卵から羽化針での発育所要lヨ数は鰍こよって輿なり．分業期でほサイロの30．8】ヨからオオの40．1！ヨまで
の幅があF），大数種ほどその期間が長くなる傾向がある。
鋤きばちの生存曲線は15穫とむに典型的なl野型な示し，卵から羽化後2−3日目までの非外役期でほ非常に死亡
率が低いが，外役活動のg享紹始とともに死亡が一定の割合で起こり，小型機ほど成虫の寿命が短い傾向な示した。また，
新女王の刹ヒの始まったコロニーでほ，働きばちの給餌対象が幼劇から獅女王成虫へと転換し．そのため幼虫の餓死
や非成虫個体に対する典食いが急用して，営巣活動のメ朗壊を促進した。
営巣j彰j間ほ穐によって興なり，短プ明償巣型のヒメほ3．5∼4ケ月に過ぎないが，長期営巣型のキイロほ7∼7．5ケ
月で，他の3郁ほキイローオオーコガターモンーヒメの順に両種の中間に位鷹する。
営巣規模ほ鋸著な稽間差があり，小娩供営巣型のヒメほ最多育房数で300房であるが，大規模営巣塑のキイロでは
10，000動こ適する。二か勇”とモンは中規模営巣彗星とみなされ戯多暫欝数で4，500房となるが，コガタは1，000厨以下で小
規模営磯遜とみなされる。
新女王の羽化期も種によって異なり．ヒメ一々ンーコガタ椚キイP−オオの順に，9月上旬より11月下旬までの幡
がある○新女王の産拇にさきがけ幼虫：働きばちの割合（‡J／W比）ほ各棟とも全営巣期を通は1牒低値に下降し，
その値は2．0前後となっている。
朗女ヨ三の産出のために必要な働きはちの個体数ほ様によって著しく輿なり，ヒメでは約20頭以」二であるが，キイp
でほ少くとも300頭以上とみなされ他種は戸！尋老の間に位俊する。
コロニ仙あたりの新女王とカースの廃山数は，山般にコーコニ…サイズが大きくなるほどその数も多くなり，その臆大
値はキイロで新女‡壬三1バ92風オス1，123風最少低ほヒメで新女王粥蘭」オス27頭であり，5種ではキイロ＞オオ＞
そン＞コガータ＞ヒメの順になっている。
廃忍冬密鍬忠，女王の亡失，天敵（オオスズメバチ，ハチクマ，スズメパチネジレバネなど），人の干吸気象，不明
などで，5穐ともに単独営巣j塁］か♭光岡営磯劇の段脚こおいて，女ヨ三の亡失による廃盤が汲も多くなりている。
鵜の年次変動に関してほ，キイロの場合，300haにおける12年間の成熟奥の数をみると3∼10巣／年と低密度の
安定型を示した。巣あたりの新女王の生産数の多さに比べ，巣の密度が非常に低いので，本機ほ強力な密度の1ヨ己調
節能力をもつとみなされるが，他の4積もその可能性をもつ。
以上の純潔から，これら5種のスズメバチほ，営賂野性，採餌寺別当三，新女王の生産紺】などの生態的地位をそれぞれ
異にすることによって棉二互の鎗争的ぎ〕済孫な少なくして，同所的に生息を可能にしているものと推激した。
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