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Entomological Science,2000,3<3):491-497

-s
Morphological Caste Differencesin the NeotropicalSwarm-
Founding PolistineWasps. IX. Polybia (Myrapetra)
ll･11IIoccidentalis
(Hymenoptera:Vespidae)
Fernando B. NoLLi*, S6ichiYAMANE2 and Ronaldo ZuccHii
iDepartamento
de Biologia,fuculdade de EloscVia Cienciase Letras de Ribeira-oPreto,
14040-90iRibeirao P}'eto,
Sao Ptiuiq Brasil
ii7･l･liLI･Ijl
igiological
LaboratorJ,,lhcutty ofEducation, Ibaraki Uhiversity,
ltdito310-8512Jopan

Abstract. Morphelogical difftrenceswere foufldbetween workers with undeveloped ovaries and an

empty spermatheca, and females with developed ovaries in two colonies of Pt)lybiaoccidentalis

collected in Ribeirao Preto,southeastern Brazil. Arnong femaleswith developed ovaries, in-

seminated femalesare designatedas queens and uninseminated ones as intermediates. Mean sizes of
most body parts of queens including alitrunk length,were largerthan those of workers. Mahalanobis
distances calculated in canonical discriminant analysis, as well as basicstatistics, indicate that
l
lI,'l///11tl
intermediates
are morphologicany closer to queens than to workers. a result rather unusual among se
far obseryed epiponines.

Key words: Vespidae, Polistinae, caste difilerences,

Pblybiaoccidentcrlis multivariate
, ana]ysis.

fiti･il/lil･llili:/
olegical differences
slight or indistinct,
and intermedi-

Introduetion atespresent (Ptirachartergus

smithii: Mateus et al.,

1997; Rseudopolybiavespiceps: Shima et al., 1998),

The study of caste differencesin social wasps has HoweveT, in some species of the group 2, queens are
recently increasedas a way to understand when and significantly smaller than workers in some characters
1111･1!11/1/111/11!E,1!llll/t1/1ill1･'vill
how pre-imaginal caste determination
has appeared in while largerin others (Richards, 1971, 1978; Shima et
thatgroup. In the two main groups of social Vespi- al., 1994), According to Jeanneet ai. (1995), Apoica
deter-
dae, i.e.,Vespinae and Polistinae,pre-imaginal pallens shews non-size-based allometry probably due
mination has been documented (O'Donnell, 1998). to an ontogenetical reprogrammation in growth
Even though caste differencesare much lessconspicu- parameters (Wheeler, 1991). Such a pattern was
6us in the Polistinae,such traitsare probablymore detected also in lipipona guerini(Huntet at., 1996)
complex than in Vespinae (Shima et al., 1994). Ac- and Jlseudopolybiadtfiicilis (Jeanne, 1996),
cording to Richards (1978)and to ouf standardized The occurrence ofintermediates, which were regard-

procedure to evaluate queen-worker differellces,

at ed as uninseminated and ovarially developed workers

leastthree main types of caste differentiation

can be by Richards& Richards (1951), adds more complex-

' categorized: 1-conspicuous size and allometric differ- ityte the panorama of the social regulation in those
i1ill..･//li,/1
present, with queens largerthan workers
ences and wasps. Because only a few behavioral observations
intermediates absent (Agetaia
pallipes and A, multi- have been made in epiponine colonies, the significance
picta: Noll et al,, 1997a; Protonectarina syJveirae:
'111/i'111
of the intermediates is practicallyunknown (butsee
Shima et ai., 1996b);2-conspicuousdimorphism pres- Forsyth, 1978 and GaFtreichet al., 1993). However,
ent, with queens smaller than workers and intermedi- Naumann's (1970) and SimOes'(pers. comm.) direct
･･Illl'ii1'1,!t1/111il/1/'il･･1/1//.:1L
ates absent Shima
(Apoica
fiavissima: et al,, 1994; observations tell that intermediatesof Protopolybia
Polybia dimidiata: Shima et al., 1996a); and 3-morph acusticutis and R exigua exigua pcrfbrm frequent ovi-
positionand oophagy. In fact, molecular studies on
* Presentadclress: Department of Entomology, The Ohio1315nell.
Brachygastra mellijica (Hastings
et al., 1998) evide-
StateUniversity,Museum of Biological Diversity,
KLnnear Rd., Columbus, OH 43212,U.S.A. E-mail: nced that the productien of males in itscoleny was

41@osu,cdu controlled by the queens even in the presence of a

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v
492 Fernande B. NoLL, S6ichiYAMANE and Ronalde ZuccH[ l-I

largenumber of intermediates. BW), and partial1ength of the forewing(WL) (see

In order te document other examples forellucidat- alsoFig, 1 in Shima et ai., 1994). The numerieal data
/:l:il//
ing the above mentioned aspects in polistine wasps, were statisticallyin relatiofl to the ovarian
analyzed

this ongoing step of our serial work deals with mor- and spermathecal states. The canonical discriminant
phological caste didefences in Polybia (Myrapetra) analysis (CDA: Rao, 1973) was perfbrmed with the
occidentalis using multivariate analyses. SAS Program Package for personal computers (ver,
6.03). For detectinginter-castesize diffbrences
for
each character, Mann-Whitney rank sum test was
Materialsand Methods Sigma Stat1.0forWindows.
perfermed using

Workers and queens analyzed here were taken from

two colenies ofI'olybia occidentaiis (Olivier)
collected
Results
in RibeiraoPreto, southeastem Brazilon September
23 and December IS, 1994 (hereaftercalled c61ony 1 Ovariandevelapmentand spermathecal contents

and colony 2, respectively) by spraying ether on the In the two analyzed samples, two extreme types of iil
nests. The colonies were taken in the daytirneand a ovaries in their development were recognized (Fig. 1):
proportion,not al1, ef returning wasps were ad- type A, with developed ovarioles bearing two to sever-
ditionallYcollected.
Colony 1 wasjust at the founding al mature oocytes, and type B, with filamentous ovari-
stage at a new nesting slte and had
yet started nest not oles bearingno visible oocyte or, ifany, rudimentary
construction. Colony 2 was collected in the worker ones, In type A females,not all the spermathecae
producing phase. A proportion ofcells had produced contained sperm. Thus, among femaleswith the type
more than one series of adults. A ovaries, those inseminatedwere regarded as queens
111ilI
In order to detect morphological differences be- and those uninseminanted were considered to be inter-
tween castes, 14 queens+intermediates(wholepopu- mediates, and the females with type B ovaries as

lation)and 100 workers (outof 353 individuals col- workers.

lected)from colony 1, and 44 queens+intermediates
(wholepopulation)and 100 workers (outof 2553 Nest and coloay conrposition
individuals collected) from colony 2 were measured Colony 1 that was right at the swarming and

under a binocular microscope. Priorto the measure- befbre the nest construction had 11 queens, 3 interme-
ments ovarial and spermathecal conditions were diatesand 353 workers (3.0% to total number of

examined for the whole individuals

to separate the females),Colony 2 (Fig.
2) had a phragmocyttarous
castes.The body partsmeasured were: head width nest with eight combs. It had 11624 cells (8076
(HW), minimum distance(IDm), gena
interorbital empty, 207 egg, 1802 larvaland 1539 pupal cells), and
width (GW), mesoscutum width (MSW), alitrunk 3S queens,9 intermediates and 2553 workers (Qltotal
length (AL),lengthof gastral tergite I (TIL),basal number of females=1.4%). The maximum number

height of Tl (TIBH), basalwidth of tergite II (T2 of meconia per cell was two, suggesting that at least
/1

/:l

E
lam B
A
.

Fig.1, Ovariesin females of Polybia occidentalis, A: Queensand intermediates; B: Werkers,

{,

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Merphological CasteDifferences
inllolybia
eccidentalis 493

llilllli!Il

Fig,2.Actiye nest ofPolybia occidentatis hanging from a tree branch, Photo taken by SidneiMateus.

Table l. Mean lengths, QIW, QII and Il'W ratios of lengths,

yalues of CANI and CAN2 on 9 characters in colony 1 of

I[l''ii'illlil///･111111il
Polybiaoceidentalis.

Means ± SD (mm)i) Ratiosi) CANI CAN2

CharactersQueens(N=11)Workers(N=100)Intermediates
(N==3)QfWQIIIIWStand-ardizedRawStand-ardizedRaw
Head
HW 2,28± O.Ol 2.25± O,02 2.28O.9SO.39
1.01*1.03*1.09* 1.01*1.021.18*
1.001.01O.92 O.16O.24O.03
6.106.33O.23
O.17 6.47
IDm O,96 ± O,Ol O.93 ± O.Ol -1.10 -29.46
'/!lli![illilli GW O.36± O.04 O.33 ± O.03 o.64 s.es
Mesosema
MSW 1.73± O,Ol 1.67± O,03 1,743.26 1.04*1.03* -e,12
O.991.001.04*1.02* -2.32 O.42-O.108.07-1.85
AL 3,27-
± O.02 3.19± O.04 O,07 1.25
Metasoma
TIL i.81± O.Ol 1.75± O.05 O,99O.971.00
1.82O,32O.92 1.03*1.031.10* 1.04*1.07111*
O,02-O,11O.30 O.04 O.63
TIAH O.31 O.02
± o.3o e.o3
± -LIO O.33 3.47-2.80
ll/Ili･ T2BW O,92 ± O.02 O.83 ± O,02 1,60 17.11 -O.26
Wing
WL 3,09 ± O,02 3,05± O,04 3.11 1.01* O.99 j.02* -e,25 -5.99O,52 IZ.65
t} Standarddeviationisnot shown for intermediatesbecauseofa small sample size.
i} Asteriskmeans a significant difference
between means for a given characteT of two castos (Mann-Whitneyrank sum test,
11'･1/
P<O,05).

two series of brood were reared there before nest of which the QIW was 1.0 (Table1). Ditfer-
ratie

collection. The real number presentmust

of workers ences werc in GW and T2BW, showing
conspicuous

be largerthan shown aboye, that is,the percentageof that queens have thicker gena and mare developed
1･////il/..I･1illl//I
queens was srrialler, becausereturning workers were gastral tergite II. Queenlintermediate ((!II)Tatios
not perfectly
collected. were equal to 1.0 in three of the nine characters, while
those forfiyecharacters were lower than 1.0and only
Morphologicaldij7?irences
betweencastes that for IDm was higherthan 1,O. But, becauseof a
･･1/t/ll/ti･
Celony 1: QueenlWorker(QfW) ratios for meas- small sarnple size of interrnediates
(N ==3),
statistical

ured characters shewed that queenswere significantly test was done between queens and intermediates.
not

larger than workers except in one character (TIAH) Finally, interrnediatelworker(I/W) ratios were
i･ki･･k

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"el.1

494 Femando B. NoLL, SaichiYAMA)"E and Ronaldo ZvccHl

II
rli
+i
'i･1
Table 2. Mean lengths, QIW, QA and IIW ratios of lengths,values of CANI and CAN2 on 9 characters in colony 2 of

folybiaeccidentalix
,1
Means ± SD(mm) Ratiosi) CANI CAN2 'i1'i

Intermediates 1/11
CharactersQueensCN-3S)Werkers
(N-leO) (N-9)Q,fWQIIVWStand-ardizedRawStand-ardizedRaw
Head
-020 -6,eg -e.09 -2.59
HW 2,33± O,07 2.24± O.06 2,36± O.02 1,otl*1,08*Lll*
O.99O.98O.9S
1.0S*LIO*1,17* 1
1ililIl'

IDm O.99 ± O.04 O.92 ± O.03 1,Ol± O.Ol 1.40 26.01 -1.16 -22.08
GW o.3g ± o.o3 o.3s± e.o2 o,41± o.o2 O.30 3.65 O.17 2.oo

Mesosoma
MSW 1.75± O.e8 ± O.06 ± O.03 1.06*1.08*O.99O.96 -O.08
1.07*1.13* -L74 -L35 -2B.23
AL 3.34± O.l91.6S 3,48± P.08
3.08± O.131.77 O.49 7.55 O.S8 B.87
Metasoma
TIL L83 ± O.12 1,70± O.10 196 ± O,07 1.08*1.07*1.12*
O.93*O.97O.96 -O.30
LIS*Lll*1.17* -4,14 1.40O.D9O.03
IS.99 :11il

TIAH o.3o ± e.o2 o.2s ± o.o2 o.31 ± o,ol O.]3 4,29 1.22o,co
T2BW O.91 ± O.07 O.81 ± O,04 O,9S± O,04 O,76 8.86
Wing
WL 3,e8 ± O,12 2.98± O.10 3.l5 ± O.05 1.03* O.98 1.06* -1.07 ･-27.2S O.74 IS,S8
i)
Asterisk means a signlficant dif[erence between means for a givencharacter of two castes (Mann-Whitneyrank sllm test,
p<c.os),3
321

21NkDU"t-2-3

Z,O
-1-2-3-4
/11･1111

-4.-2oCANI2 4 6 -s.e-3.3-1.6 O.1 1.e 3.5B

A CANI
lili
Fig. 3. Discrimination among queens (Q),intermediates (I)and workers (W) ofPolybia occidentatis in colonies 1 (A) and
2 (B),respectively, based on the canonical discrirninant
analysis using 9 metric characters.

'
higher than l.O in all These
characters (-2 to 2), while intermediates had larger
analyzed. workers
/
thatqueens are distinctly
results indicate largerthan values of CAN2 (Oto 2.5)than queens (-2 to 1) and
workers, and intermediates
are moTe similar to queens workers (-2 to 2.S).
than to workers. Such differences can be yisuallzed For determining CANI (Table1), HW, IDm, WL
based on the Tatios of alitrunk length <Qrw=1.03, and, especially, T2BW were the most ifnpeTtantchar- '
QII==1.0,IIW=1.02),whichcanbeareferencelength acters. MSW, WL, GW and, especially, IDm were
size difference(Shimaet
to evaluate the inter-caste al,, most important among the characters examined for
1994). determiningCAN2. The Mahalanebisdistances(D2:
Plots of each caste against the firstand second Anderson, 196S) calculated through the CDA were
canonical variables (CANI and CAN2) calculated on 18,Obetweenqueens and workers, 3.1betweenqueens
nine morphological characters showed an overlap of and intermediates and, 17.4 between intermediates
queens intermediates,
and and clear
of separation and workers. UsingD2 as x2 values (Rao,1973), QfW
these ovarially developed individuals(queens and and IIW
distances calculated were highly significant
intermediates) from workers (Fig, 3, A). Queen$and (P<O.OOI),
while QII distances were not significant
intermediates had larger values of CAN1 (3to 5) than (P<O.20).

ttt

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Morphological inR)lybiaoccidentalis
Caste Differences 495

i Colony 2: Q'fW and IIW ratios were sigriificantly do,

'analysis
higherthan 1,Oin all characters analyzed (Table 2). Canonicaldiscriminant also revealed such

As in colony
tll･f,･ll[ 1, differences between queens and work- differences. Comparing the two colonies, females in
ers were conspicuous in GW and T2BW, in which colony 1 presentedlargerMahalanobis distances(D2)
1.10. But, in most characters, QfW than those in colony 2, but,in some way, caste differ-
ratios exceeded
ratios were higher than those in colony 1, In QII ences fo11owedthe same pattern in both cases. In

ratios all character$ were lower than 1,O showing a addition, in both colonies ovaTially developed but
el
tendency that queens are smaller than intermediates, uninseminated fernales (intermediates) were closer to

but the diffbrence

was significant only in TlL. queens than workers.

TheCDA based on nine characters showed

result of Queen-worker differences of P: occidentaiis were,

however, less conspicuous POlybia

patternobserved in colony 1,that is,queens as compared with
a similar
1 overlapped with intermediates but these two diffbred (lldZ ) scutellaris which presents well pronounced mor-

in size from workers (Fig. 3, B). However, the values pho-physiological caste differentiation (Nell al.,et
I{ of CAN1 indicated that the differencesof queens and 1997b). Sharp distinctness in the ovary conditions
intermediates from workers were less distinct if com- between queens and workers occurs with significantly
larger D2 in R (D2=62.8), Ageiaia
paredwith those in colony 1 (O,1 to 3,5, 1.8 to 3,O and vicina
IIIIllltlljl･iliLellllli'1i'li'il[tIl.l.li!/ul/IIe1'1ill/tllt/111i scutellaris

-3.3 to O.1,respectively), On the other hand, inter- (207.2, Baio et al., 1998),A, paUipes and A. multipicta

mediates showed higher values of the second canoni- (124.7 and 111.0,respectively: Noll et aL, 1997a).
Species of D2:
cal variable (-O.5to 3) than queens and workers of other genera show smaller values

Rseudopolybia (1.0: Shima et aL, 1998),

(-3.5to 3). vespiceps

For determiningCANI (Table 2),AL, T2BW, WL Apoica Y7avissima (11,3:Shima et at., 1994), Pblybia
and especially IDm were most important among char- (dytindroeca) dimidiata (6.8: Shima et al., 1996a),
acters analyzed, WL, IDm, MSW and, especially, TIL Protopolybia exigua (5.0: Noll et al., 1996), Rarachart-

were the most importantcharacters for determining ergus smithii (10.3: Mateus et al., 1997) and Protone-
CAN2. TheD2 was 12.6between queens and workers, ctarina sylveirae (8.6: Shimaet at,, 1996b).Except for
2,4 between queens and intermediates and, 11.2 the occurrence of intermediates, pattern of caste
between intermediates and workers. These values also differentiation in R occidentalis is closely similar to

indicateslight separation betweenqueens and interme- that observed in Protonectarina sylveirae(Shima et

diateand clear distinction between ovarially devel- al,, 1996b).

oped individuals and workers. Using Di as x2 values An importantproblem to be solved refers to the role
the intermediates, because they may provide a clue
(Rao,1973), QfW and I/W distances calculated were of

highlysignificant (P<O,Ol) while QII were insignifi- to the sociogenesis of epiponine colonies. Intermedi-
as representing a ceftain phase of work-
cant (P>O,1). ates, regarded

ers which developed their ovaries (Richards, 1978;

Simaes, 1977), have been recorded in species belong-
Discussion ingto seyeral genera: Dotopolybiaacutiscustis (cited as

As preyiouslysuggested by Richards (1978), the R pumila. Naumann, 1970),R exigua exigua (Noll
1･l[tt/(llt present results confirm that Polybia (Myrqpetra) oc- et al., 1996), Polybia (T}'ichothorax) chtysothorax,

cidentalis presents biometrically clear caste differences R (Apopolybia)jurinei (Richards & Richards, 195l),
developed females,whether in- Parachartegus smithii (Mateusetal., 1997) and Ilseud-
between oyarially
et al,, 1998),Pbrachartergus
seminated or not, and workers. In both colonies, opolybia vespiceps (Shima
queenswere larger than workers in all measured char- fraternus and Angiqpolybia spp. (Richards & Rich-

and thisspecies belengsto the firsttype ofcaste

/liil･il･ilttllli･liI11illl/
acters, ards, 1951), Chartergellus communis (Itichards &
differentiation. Among characters, gena widthi and Richards, 1951; Mateus et al., 1996),Brachygastra
basal width of gastraltergiteII had Q!W ratios con- scuteilaris (Carpenter & Ross, 1984), B. Iecheguana
spicuously higherthan in ether characters.
The fact (Machado et al., 1988) and B, mettijica (Hastings
that queens had gastral segments much larger than et al,, 1998).
workers is consistent with other so far observed spe- In the presentcase, however, queensand intermedi-
cies, but the possession of gena thicker than that of ates were only slightly separated by CAN2 values,

species (Shima This result resembles the case observed in Agelaia

workers is inconsistent with other
et al., 1994, 1996, 1998, ctc,). It is open te further vicina (Sakagami et al,, 1996; Baio et al., 1998) in
queen-sizedbut uninseminated femaleswere
studies why queens have thicker gena than workers which

itt
L
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496 Femando
!l
iLi
B,NoLL. S6ichi YAMANE and Ronaldo ZuccHI

found. It suggests that unisemlnated but ovarially Hasimgs, M. D., Queller, D. C., Eischen,F. & Strassmam, J.
deyeloped femalesfound in I]Olybiaoccidentalis can be E. 1998, Kin selectien, relatedness and worker control of
young queens as regarded in other species by Richards
reprodllction in a large-colony epiponine wasp, Brachy-
gastra melldica, Behavioral EbolQgy, 9: 573-581.
(1978) and Gastreich et al. (1993). Chandrashekara Hunt, J. H,, Schmidt,D. K., Mulkey, S.S, & Williams,M. A.
& Gadaghar (199l)considered that femalesofRopa- 1996. Caste dimerphism in thipenaguerini /
tidia marginata (Hymeno-
eclose with different reproductive ptera: Vespidae): Further eyidence for larvaldetermina-
potentials, and that perharps all eclosing femalesare tion.JeurnalofKansas Entomolagical Society, 69/362-369. i1lt
not potentiaily capable of becoming egg layers.Such Jearme, R, L, 1996. Non-altometric queeri-worker dimorphism
an idea may be also applied to some epiponines like in}lseudopolybia (Hymenoptera:
dCfiiciiis Vespidae),.lour-
nal ofKansas Entomoiegical Seciety, 69: 370-374.
A. vicina and A occidentalis, in which interTnediates
Jeanne,R. L., GraC C. A, & Yandell, B. S. 1995. Non-size-
are morphologically to queens than to workers.
cleser
basedmorphelogical castes in a social insect,
IVlrturk,issen-
The intetmediates ofthese speoies could be interpreted schtijlen, 82: 2P6-298. 1]i
as umnseminated queens with a low reproeluctive Machado, V. L.,Gravena, S. & Giannotti,E. 19S8. Analise
potential and the contribution to male production can populacional e morfometrica em uma colonia de Brachy-
be considered, An alternative explanation isthat un- gastratecheguana(Latreille, 1824) na fase reprodutiya.
fertilizedeggs Anais da Sbciedade Entomol6gieado Btvsil, 17: 491-506.
could be used forfoed as observed in
Mateus,S., Noll, F. B. & Zucchi,R. 1997. Morphologicalcaste ttiil
Protopolybia exigua (SimOes,1977) and P. acutiscutis dtfferencesin Neotrepical swarrn-founding polistine
wasps:
(Naumann,1970), Their role in social regulation in llarachartetgtts smithii <Hymeneptera: Yespidae). Jburnal
the colQny, however, remains speculative. oftheNtnv Ybrk EntomolagicalSbciety, le5: 129-139.
Mateus, S., Noll, F, B. & Zucchi,R. 1996. Morphological caste
diff.e..r.enees.
in Cha.rtergR.llu.s
communis (Hymenoptera,
Acknowledgments Yespidae. Epiponini).Proceedings ofXIth International
The authors acknowledge the financial support by
thngress of
Entomolqgy.p. 409.
Naumann, M.G. 1970. The nesting behavior ofPtotopolybia
FAPESP (Fundagtto de Amparo a Pesquisado Estado
pumila in Panama (Hymenoptera, Vespidae). Ph.D.

lI
de Sao Paulo) and CNPq (Conselho Nacional de thesis,
University of Kansas.
Desenvolvimento Cientifico e Tecno16gico), This Noll,F. B.,Mateus,S. & Zucchi, R. 1996. Morphologicalcaste
study was in part supported by a grant-in-aid (No. difierencesin Neotropical swarm-founding pelistine
wasps.

10041154) from the Japan Ministry of Education, V. Protopolybia exigtta exigua (Hymenoptera: Vespidae).
Science, Sportsand Culture. Specialthanks are given JburnaloftVbwVbrk EntomologicalSbciety, 104t 61-68,
to Sidnei Mateus for histechnical assistance and for a
Noll,F. B.,Simbes,D. & Zucchi, R, 1997a.Morphelogiealcaste
differencesin Neotropicalswarm-founding pokstinewasps.
photo of nest at our disposal. VII. Ageiaia rn. multiptcta and A, p. pallipes (Hymeno- l1/
ptera:Vespidae). EthoJogy,Ecotogy & Evolution, 9: 361-
372.Noll,
References F. B., Mateus, S. & Zucchi,R. 1997b. Morphological
Anderson, T. W. 1968. Introductionto multivariate statistical easte diffbrencesin the Neotropicalswarm-founding and
/tl

analysis, John Wiley and Sons Inc., New York. polygynous polLstinewasps, VIII, folybia scuteltaris. Stud-
Baio M. V., Noll,F. B.,Zucehi,R. & SimOes,D. 1998. Non- ieson the IVbotropical Flruna and Environment, 32: 76-80.
allometric caste differences in Agelaia vicina (Hymeno- O'Donnell, S. 1998, Reproductivecaste deterrnination in eu-
ptera, Vespidae,Epiponini).Sociobiotogy, 34: 465L476. social wasps (HyTnenoptcra: Vespidae).Annuat Review of
Blackith,R, E. 19SS. An analysis of polymorphism in social EntomolQgy,43: ]23-346.
wasps. Insectes Sbciadeh, 5:263-272. Rae, C, R, 1973. Linearstatistieat infefence. John WMey and
Carpenter,J,M. & Ross, K. G, I9S4. Colony composition in Sons,New York,
four species of Polistinae from Suriname, with a deserip Richards,O. W. 1971. The biologyof the social wasps (Hy-
tion ofthe laTva ofBrachJtgastra scutellaris (Hymenoptera, inenoptera: Vespidae).BiologicatReview, 46:4B3-52S.
Vespidae).Rsp,che, 91: 237-250. Richards, O.W. 197g. The social wasps of the Americas
Chandrashekara, K. & Gadagkar, R. 1991. Unmated
queen in excluding the Vespinae. British Museum (Natura] Histo-
the primitively eusocial wasp Ropalidiamarginata !,ep. ry), London.
(Hymenoptera, Vespidae). Insectes Sbciaux,3S: 213-216. Richards,O. W. & Richards,M. J, 1951. 0bservatiens on the
Forsyth,A. 1978. Studies en the behavioralecology of poly- secial wasps efSouth America. 7)'ansaction qf'theEntomol-
gynous social wasps. Ph, D. dLssertatien, Harvard Univer- ogical SocietyofLondon,102: 1-170.
sity, Cambridge. Sakagaini, S,F., Zucchi, R.. Yamane, S6., Noll, F,B, &
Gastreich,K, R., Strassrnann, J,E. & Quetter, D. C. 1993. Camargo, J, M. F. 1996. Morpholegicaleaste difflerences
Detetminants of high genetic relatedness in thc swarm- inAgelaiavicina,theNeotropiealswarm-foundingpolistine
founding wasp, Ihotopolybia exigua. Etholqg}t Ecology & wasp with the largestcolony s{ze among social wasps
Evolntien, 5: 529-539, Vespidae).Sociobiotogy,
(Hymenoptera: 28: 1-17.

'L

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The Entomological Society ofof Japan
Japan

lliii't., MorphologicalCasteDifibrences inilolybiaoccidentalis 497

Shima, S.N., Yamane, SO, & Zucchi,R. 1994. Morphological Morphological caste differences in the Neotropicalswarm-
caste differences in some Neotropicat swam-founding foundingpolistinewasps. IV. Rsettdopolybia vespieeps, with

Yes- on the role of intermediate

polistine wasps. I. Apeicanavissima (Hymenoptera, preliminary censiderations

femalesin seeial organization of the Epiponini (Hymeno-

pidae),Jqpanese Jburnal ofEntomolQgp, 62:811-822.
Shima,S. N., Yamane, Sa. & Zucchi,R, 1996a. Morphological ptera, Vespidae). Jburnal ofHymenoptera Resee rt]h, 7: 280-
caste diferencesin some Neotropical swarm-founding 295.Simaes,
D. 1977. Etologia e diferenciagao de casta errt alguTnas
polistinewasps. II. Ilolybia dimidiata (Hymenoptera,
Yespidae). JapaneseJburnalofEntomology,64: 131-144. vespas sociais (Hymenoptera, Vespidae). Ph, D, thesis,
Shima, S.N,, Yamane, SO. & Zucchi,R. 1996b. Morphologi- RibeiraQPreto-USP.
l'i

nb,
cal caste differencesin somc Neotropicalswarm-founding Wheeler,D. E. 1991, 1[hedeyclopmerttal basis of worker caste

pelistillewasps. III. Protonectarina sylveirue (Hymeno- pelymorphism in a"ts. American IVbturatist, l38: 12!8-
ptera, Vespidae). Bttlletin of the Fhculty of Education, 1238.
JbarakiUhiversity (IVtrturalStience), 45: S7-6T,
Shima, S.N., Noll, F. B., Zuechi,R. & Yamane, SO. 1998, (Received January 4, 2000; Aceepted June 20, 2000)
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