Pacific Science (1995), vol. 49, no. 1: 42-54 © 1995 by University of Hawai'i Press. All rights reserved

Variation in Reproductive Strategy of the Tropical , Ropalidiafasciata (: ), in Okinawa in Relation to Island Environmental Conditions 1

YOSIAKI ITo 2

ABSTRACT: The tropical paper wasp, (Icariola) fasciata (F.), nests on leaves of gramineous plants ( Andus. and sugarcane) in Okinawa, where there are frequent, strong typhoons. In Taiwan and Java, where the effects of typhoons are less severe, most nests are on tree twigs. A similar difference is seen in nests of Ropalidia (I) marginata (Lepeletier) in the Northern Mariana Islands and in India. Okinawan R. fasciata also exhibits quite flexible social behavior, low frequency of intranidal dominance behavior, construction of satellite and multiple-comb nests, absconding swarming, and initiation and development of nests in late autumn. Study of the divergence of social habits in eusocial wasps on Pacific Islands will enhance our under­ standing of social evolution in .

THE SUBFAMILY ofthe family Ves­ tively eusocial groups without passing pidae occupies a unique position in the evo­ through a monogynous stage. lution of , because this On the other hand, Carpenter (1991) pos­ group has both primitively eusocial (without tulated that short-term monogyny among distinct queen-worker polymorphism) and multiple foundresses is the primitive state in highly eusocial (with distinct polymorphism) the Polistinae, and single-queen and multi­ . The latter are characterized by a so­ queen social systems are both derived sepa­ cial system with multiple queens (that is, they rately from this ground plan. Carpenter's ar­ are "permanently polygynous" [Jeanne gument was based on the fact that initial 1991)). This polygynous social structure is polygyny (multi-female founding of new remarkably different from that ofhoney bees, nests) of primitively eusocial (e.g., stingless bees, and hornets (except for a few Pardi 1942, 1946, West-Eberhard 1969), tropical species [Matsuura and Yamane (Jeanne 1972), and Ropalidia 1990)), in which each colony has a single (Gadagkar and Joshi 1983) changed to func­ queen (Wilson 1971). tional monogyny through dominance be­ There is a debate about the evolution of havior, where only the top-ranked females such multi-queen social systems, which might could leave reproductive progeny. Carpenter reduce relatedness among nestmates (Jeanne defined this as "short-term monogyny." If 1991, Spradbery 1991, Ito 1993a,b). Thus, primitively eusocial species are found in the role of kin selection (Hamilton 1964) in which many colonies retain multiple egg-lay­ evolution of eusociality might be limited. ers, the possibility of evolution of multi­ West-Eberhard (1978) proposed the polyg­ queen systems from such species, and the ynous family hypothesis, which states that polygynous family hypothesis, will have polygynous colonies of these polistine genera more support than at present. could have evolved from polygynous, primi-

Ropalidiafasciata IN JAVA AND OKINAWA I Manuscript accepted 27 April 1994. 2University of Okinawa, 555 Kokuba, Naha City, The genus Ropalidia is a useful subject to Okinawa, Japan 902. investigate the evolutionary process in euso- 42

¥ , ±J4es.,,\ ffl\9ff, WiHWdlftM #!!fltJJlfilhM WMWHi1i"EM'SMif 'HFt9iRW Reproductive Strategy of -1T6 43 cial species. This large genus (136 described nests also tended to establish their nests again species found in Africa, South Asia, Aus­ on trees (unpubl. data). tralia, and Oceania) has both higWy and Turillazzi and Turillazzi (1985) observed primitively eusocial species and both monog­ the social behavior of R. fasciata in Java and ynous and long-term polygynous (multi­ found that, in each of two postemergence queen) species (Gadagkar 1991, Jeanne 1991, nests (i.e., nests after emergence of progeny), Ito 1993a). The subgenus Icarielia includes only dominant females had developed ova­ higWy eusocial, multi-queen species; the ries. other four subgenera, including the largest In a pre-emergence nest studied by Tur­ subgenus, Icariola, are all primitively euso­ illazzi and Turillazzi (1985), the status of a cial except for R. (Icariola) ignobilis (Saus­ dominant foundress was later usurped by the sure) (Ito 1993b). Ropalidia also includes in­ first progeny, whose status was usurped in dependent-founding and swarm-founding tum by her younger sister. The dominant fe­ species (Table 1). male did not perform extranidal work. Only Ropalidia (Icariola) fasciata (F.) occurs dominant females can initiate food-request­ from India to Taiwan and Indonesia; the ing behavior toward returning foragers northern limit is the Ryukyu Islands. In Tai­ ("kiss" [Ito 1983]). Thus, in Indonesia, multi­ wan, this species constructs nests that hang foundress colonies changed to functionally from twigs of trees (So. Yamane, pers. monogynous ones through dominance hier­ comm.). In Indonesia, nests are often hung archy. from twigs of trees and occasionally from Although only four R. fasciata nests were leaves of grasses (So. Yamane, pers. comm.). observed by Turillazzi and Turillazzi (1985) In Okinawa, in the Ryukyu Islands, nests of in Java, this situation is similar to those of R. this species are almost always constructed (Icariola) marginata (Lepeletier) in India hanging from leaves of a grass, Miscanthus (Gadagkar 1980, Gadagkar and Joshi 1983) sinensis Andus., or sugarcane (Ito 1983). Al­ and R. (Icariola) variegata (Smith) in Suma­ though nests have been seen rarely on twigs tra (So. Yamane 1986), in which the number of small trees, these were probably estab­ of egg-layers was limited to one per nest ex­ lished by a rare, genetically different race, cept in very large nests. because females that emerged from these In Okinawa, 55% of R. fasciata nests

TABLE I SUBGENERA OF THE GENUS Ropalidia

SUBGENUS SPECIES TYPE OF FOUNDING TYPE OF NEST

Primitivelyeusocial lcariola Most species· Independent-founding Arboreal nest Some speciesb,' Swarm-founding? Nonenveloped nest in cavities Ropalidia maculiventris GUerinb,d Nonenveloped nest in cavities Polistratus bambusae Richardsd Swarm-founding? Nonenveloped nest in cavities Anthreneida Some species' Independent-founding Arboreal nest sumatrae (Weberl,J Swarm-founding? Nonenveloped nest in cavities Highly eusocial lcarielia All speciesb,. Swarm-founding Enveloped nest

Q R. marginata (Gadagkar and Joshi 1983), R. fasciata (Ito 1983), R. revolutionalis (Saussure) (Ito 1987). • Construct large colonies (> 1,000 adults) including many egg-layers. , R. mackayensis Richards and R. trichphtalma Richards (Jeanne 1972), R. socialistica (Saussure) (pers. obs.). d Spradbery and Kojima (1989). , R. taiwana koshunensis Sonan (Iwata 1969). f Aramaki (1985). • R. montana Carl (Yamane et al. 1983), R. romandi (Saussure) (S-N. Shima et aI., unpubl. data), R. flavopicta (Smith) (So. Yamane and K. Ikawa, unpubl. data). FIGURE I. Foundation of a nest by a group of foundresses of Ropalidiafasciata (Okinawa, 1983).

eN 56' it B! i , S&W%iJ&&&& Reproductive Strategy of Ropalidiafasciata-ITo 45

TABLE 2

FREQUENCY OF DOMINANCE-AGGRESSIVE ACTS AMONG CO-FOUNDRESSES OF SOME POUSTINE WASPS (PRE-EMERGENCE PERIOD)

NO. OF COEFFICIENT OF a SPECIES LOCALITY n DOMINANCE ACTS VARIATION AUTHORS

Polistes dominulus Italy 1 1.95 Pardi 1942 canadensis (L.) Panama 6 1.36 ± 0.55 40 Ito 1985a Ropalidia fasciata Okinawa 30 0.89 ± 0.70 89 Ito 1993a gregaria (Saussure) Darwin, Australia 4 2.85 ± 0.86 30 Ito and Yamane 1992

a Number ofobservations of more than 30 min.

(n = 251) were established by foundress What causes the low frequency of dom­ groups of two to 22 individuals (Figure 1), inance-aggressive acts and effective oviposi­ but the frequency of dominance-aggressive tion by subordinates? The survival rate of behavior among co-foundresses was signifi­ nests founded by single females was far lower cantly lower than in some Polistes (P < 0.05, than that of nests founded by multiple fe­ Mann-Whitney two-tailed U test) (Table 2). males. When a nest was destroyed by ty­ In Okinawa, I found the following: (1) not phoons or predation by ants, the multi-foun­ only dominant but also subordinate females dress colonies reconstructed their nests, but could initiate a "kiss" toward dominant and the single-foundress colonies did not, even subordinate females (Ito 1983, 1985a); (2) co­ after emergence of some progeny (Table 4). foundresses coexisted on a nest until July or Thus, the number of cells constructed per August, 2-3 months after the emergence of foundress (which was considered to be an in­ progeny females (Figure 2); (3) several fe­ dex of fitness of the foundress) was signi­ males laid their eggs on these nests; and (4) at ficantly larger in nests established by six to 10 least some eggs laid by subordinates survived foundresses than in single-female nests, be­ (Ito 1985b, 1993a). cause of the high survival rate of multi-fe­ As shown in Table 3, in R. fasciata even male nests (P < 0.05, Mann-Whitney two­ females whose status was the third rank or tailed U test) (Figure 3). lower can oviposit and eat eggs laid by other Okinawa is famous for its frequent, strong females. This situation is quite different from typhoons. This may be a reason that these that in Polistes dominulus Christ (pardi 1942, wasps construct their nests on grasses (grass 1946, Gervet 1964) and P. fuscatus (F.) leaves are soft and nests on these leaves are (West-Eberhard 1969), where dominant fe­ better able to survive typhoons than nests on males lay significantly more eggs and eat eggs twigs) and for the relatively peaceful coex­ laid by subordinates at a significantly higher istence of many foundresses. rate, and, among subordinates, only second­ Other characters of behavioral flexibility rank foundresses oviposit and perform 00­ of Okinawan R. fasciata have been observed. phagy; third- or lower ranked females never Colonies often established nests with multiple perform these actions (P < 0.05, Fisher exact combs and satellite nests (Ito 1986). The dif­ probability one-tail test) (Table 3). ference between multiple-comb nests and Although monopolization of effective ovi­ satellite nests is that in multiple-comb nests position has sometimes been observed in R. every comb was used by all colony members fasciata (Kojima 1984, Iwahashi 1989), this is until the end ofthe colony cycle; in the latter, not a rule. Thus, the social behavior of the original nests were abandoned after estab­ Okinawan population of R. fasciata is char­ lishment of satellite nests. acterized by its great flexibility. In addition, Okinawan R. fasciata col- FIGURE 2. At least three Ropalidia fasciata co-foundresses (marked by arrows) were surviving on their nest 3 months after emergence offirst progeny (Okinawa, 1991).

IWJliULJiiUCJ££ZEUAiS¥ U• 'E' ,I ffidiSi&iJ !;a Iii I'Wb fU!!!WU iilbiHMLiMW:ie)t.!!Ji§&£g Ii' TABLE 3

OVIPOSmON AND OOPHAGY BY DOMINANT AND SUBORDINATE FEMALES IN SOME POLISTINE WASP CoLONIES

OVIPOSmON BY OOPHAGY BY

DOMINANT SUBORDINATE % BY DOMINANT DOMINANT SUBORDINATE % BY DOMINANT SPECIES STAGE" n FEMALES FEMALES· FEMALES FEMALES FEMALES FEMALES

Ropalidia fasciata' Pre- 22 3 4 (4) 43 0 1 0 Post- 22 11 10 (8) 52 1 6 14 Total 44 14 14 (12) 50 1 7 13 Polistes dominulus Pre-d 2 55 33" 59 31 7" 82 PostJ 2 131 69" 66 164 36 82 Total 4 186 102" 65 195 43" 82 Post-g 41 6" 87 Polistes fuscatus Pre_h 1 9 5" 64 4 2" 87

"Pre-, pre-emergence stage; post-, postemergence stage. • Numbers in parentheses are the mean number of eggs surviving.until the end of the observation period. 'Ito (1993a). ·Pardi (1942). "By second-rank females only. I Pardi (1946). • Gervet (1964). 'West-Eberhard (1969). 48 PACIFIC SCIENCE, Volume 49, January 1995

TABLE 4 SURVIVAL RATE OF SINGLE-FEMALE AND MULTI-FEMALE COLONIES AND NUMBER OF NESTS RECONSTRUCTED AFTER DESTRUCTION OR ABANDONMENT OF ORIGINAL NESTS IN THE OKINAWAN POPULATION OF Ropalidiafasciata

SURVIVAL RATE OF NO. NESTS RECONSTRUCTED BY

% OF MULTI- SINGLE-FEMALE MULTI-FEMALE SINGLE-FEMALE MULTI-FEMALE YEAR FEMALE COLONIES COLONY COLONY COLONY COLONY UNKNOWN

1983 53.4 (58) 46.2 (26)* 86.7 (28) 0 4 1 1984 52.9 (119) 36.4 (55)* 77.8 (63) 1 3 2 1985 63.8 (47) 33.3 (15)* 88.0 (25) 0 8 9 1986 51.9 (27) 30.8 (13)* 76.9 (13) 0 1 0 Total 55.0 (251) 38.5 (109)* 81.9 (129) 1 16** 12

NOTE: Numbers in parentheses denote sample size; values for survival rate are not the same as values for percentage of multi­ female colonies because some nests were sacrificed for dissection. *Difference between values ofsingle-female and multi-female colonies was significant (P < 0.00I, X2 test). ** Difference between values of single-female and multi-female colonies was significant (P< 0.001, Fisher exact probability test).

Until emergence of progeny Until middle or late July 100 22(7) 10(4)

cu .~ 50 :J .....(J) (J) CD Z

OL- -L-=-_~ _ (J) (J) ~ "D C :J .2 t; 10 en CD 'ij5 C 151 _u=n:l o C ~ .- CD .D E 0'---'---'------L..--__---.l_---.l...-l...._---l L-- --L_ :J 11 + 1 2-5 6-10 11 + Z Initial number of foundress

FIGURE 3. Productivity (number of nest cells constructed per foundress) and survival rate of single-female (one initial foundress) and multi-female colonies of Ropalidia fasciata. The numbers in the top graphs show the mean number of nests studied and, in parentheses, the number of observed cases (product of number of survey stations and years). The numbers in the bottom graphs show total number of foundresses at the initial stage of nest founda­ tion (Okinawa, 1983-1985). Vertical lines indicate SEM. onies often abandoned their original nests This is nest relocation or absconding swarm­ without any sign of predator attack or me­ ing as defined by Jeanne (1991), first reported chanical damage and constructed new nests for primitively eusocial wasps. (Figure 4). Before the new construction, cell I also found that R fasciata colonies contents of the original nests were removed. sometimes reconstructed their nests even in

'd#eM"teeew"'" icEiEe"uFSM* &' 1M FIGURE 4. Nest relocation (absconding swarming [Jeanne 1991)) in Ropalidiafasciata. A, Original nest was aban­ doned after removal of cell contents (26 July). B, A group of females constructing a new nest (arrow) 50 cm above the original nest (26 July). C, The same group of females and nest (arrow) on 29 July (Okinawa, 1991). 50 PACIFIC SCIENCE, Volume 49, January 1995

FIGURE 5. Reconstruction of a nest by progeny groups of Ropalidiafasciata in autumn (21 August 1992, Oki­ nawa). No foundress was seen, but this nest produced many female progeny. autumn (although more than 90% of the col­ pupae, but also larvae and eggs were seen. onies complete their development in October This nest was still active and growing in Jan­ and November). Figure 5 shows a nest being uary (Y. Sadoyama, pers. comm.). Dissection reconstructed by a large group of progeny at of new females taken in late November from the end of August. three nests established in autumn showed Figure 6 shows a nest established in late that all of them were inseminated and 84% November. Even at this late date, not only had mature ovaries. This is quite different

Hi 1M &&Mi$ -;;*;4&*35 ; Reproductive Strategy of Ropalidiafasciata-lT6 51

FIGURE 6. A Ropalidiafasciata nest established by four females in late November. Pupae, larvae, and eggs were seen. This nest developed until January (Okinawa, 1992-1993).

from behavior of any known Polistes in tem­ NEED FOR STUDIES OF SOCIAL WASPS perate and subtropical areas (e.g., Turillazzi ON ISLANDS 1980), in which females that emerge during autumn are inseminated but do not develop As mentioned before, R marginata shows ovaries before overwintering. functional monogyny in India (Gadagkar These facts indicate that social behavior of 1980, 1991, Gadagkar and Joshi 1983). This Okinawan R. fasciata is quite flexible, possi­ species occurs in a wide range from India to bly because of special habitat conditions on the Northern Mariana Islands and Iwo lima islands, especially strong predation pressure of the Ogasawara (Volcano) Islands. In the by ants and frequent typhoons. Relocation Northern Mariana Islands, where typhoons of nests by groups in R. fasciata is con­ are common, nests are often constructed sidered to be an intennediate stage between hanging from leaves of small herbal ferns independent-founding of many Icariola and (Figure 7; S. Miyano, pers. corom.). In India, swann-founding of known Icarielia. This however, nests are found under eaves or on suggests that the multi-queen social system of tree twigs (Figure 7). I speculate that there the subgenus Icarielia could have evolved may be differences between the social be­ through swann-founding, primitively euso­ haviors of Marianan and Indian populations. cial, polygynous species, as in West-Eber­ Two interesting species of Ropalidia have hard's polygynous family hypothesis. recently been found in Madagascar. One of 52 PACIFIC SCIENCE, Volume 49, January 1995

FIGURE 7. Left, A nest hanging from eave of a house (Bangalore, India, 1989). Right, A R. marginata nest hanging from an herbal fern on Anatahan Island, Northern Mariana Islands (July 1992; photograph by S. Miyano). them, R (Icariola?) formosa (Saussure), Several species of another social wasp nests gregariously, often in dense clusters of genus, North American Polistes, have been hundreds of nests under overhanging rocks introduced into Hawai'i. The extent to which (Wenzel 1987), as R. (Icariola) plebeiana their behaviors have differentiated from those Richards does in Australia (Ito et al. 1988). of mainland populations is an interesting Most nests had only one adult female (mean subject for future study. = 1.03 females, n = 262) and only 10 to 20 cells (although the largest nest had 56 cells). Wenzel (1987) concluded that this is a nearly LITERATURE CITED solitary species. The other species, R. (Icar­ iola) ignobilis, had morphologically distinct ARAMAKI, H. 1985. Female dimorphism in castes despite its small colony size (Wenzel the tropical paper wasp, Ropalidia suma­ 1992). This is unique because all known trae. B. Ed. thesis, Ibaraki University, higWy eusocial polistine species construct Mito, Japan. large nests with hundreds of adults. Although CARPENTER, J. M. 1991. Phylogenetic rela­ Madagascar is far larger than Pacific Islands, tionships and the origin of social behavior founder effects and lack of competitors may in the Vespidae. Pages 7-32 in K. G. Ross affect the evolution and maintenance of such and R. W. Matthews, eds. The social unique social characters. biology of wasps. Comstock Publication

ffi i#!ffi1MUlUUfh 1i W5B e "1'''&6;0''. &9A4 Reproductive Strategy of Ropalidia fasciata-ITO 53

Associates, Cornell University Press, pothesis. Oxford University Press, Oxford, Ithaca, New York. U.K. GADAGKAR, R. 1980. Dominance hierarchy ---. 1993b. The evolution of polygyny in and division of labour in the social wasp, primitively eusocial polistine wasps with Ropalidia marginata (Lep.) (Hyme­ special reference to the genus Ropalidia. noptera: Vespidae). Curro Sci. (Bangalore) Pages 171-187 in L. Keller, ed. Queen 49: 772-775. number and sociality in insects. Oxford ---. 1991. Be/onogaster, Mischocyttarus, University Press, Oxford. Parapo/ybia and independent founding ITO, Y., and So. YAMANE. 1992. Social be­ Ropalidia. Pages 149-190 in K. G. Ross havior of two primitively eusocial wasps, and R. W. Matthews, eds. The social Ropalidia sp. ill. variegata and R. gregaria biology of wasps. Comstock Publication gregaria (Hymenoptera: Vespidae) in Associates, Cornell University Press, Northern Territory, Australia, with spe­ Ithaca, New York. cial reference to task specialization and GADAGKAR, R., and N. V. Joshi. 1983. mating inhibition. J. Ethoi. 10: 63-74. Quantitative ethology of social wasps: ITO, Y, SO. YAMANE, and J. P. SPRADBERY. Time-activity budgets and caste differen­ 1988. Population consequences of huge tiation in Ropalidia marginata (Lep.) nesting aggregation of Ropa/idia p/ebeiana (Hymenoptera: Vespidae). Anim. Behav. (Hymenoptera: Vespidae). Res. Popui. 31: 26-31. Ecoi. (Kyoto) 30: 279-295. GERVET, J. 1964. Le comportement d'oopha­ IWAHAsm, O. 1989. Society of Ropalidiafas­ gie differentielle chez Polistes gallicus L. ciata. Pages 3-209 in O. Iwahashi and So. (Hymen. Vesp.). Insectes Soc. 11: 343­ Yamane, eds. Societies of Ropalidia 382. wasps. Tokaidaigaku Syuppankai, Tokyo HAMILTON, W. D. 1964. The genetical theory (in Japanese). of social behavior, I and II. J. Theor. BioI. IWATA, K. 1969. On the nidification of Ro­ 7: 1-52. palidia (Anthreneida) ·taiwana koshuensis ITO, Y 1983. Social behavior ofa subtropical Sonan in Formosa (Hymenoptera: Vespi­ paper wasp, Ropalidia fasciata (F.): Field dae). Kontyu 37: 367-372 (in Japanese observations during founding stage. J. with English summary). Ethoi. 1: 1-14. JEANNE, R. L. 1972. Social biology of the ---. 1985a. A comparison of intracolony Neotropical wasps Mischocyttarus drew­ aggressive behaviours among five species seni. Bull. Mus. Compo BioI. Harv. Univ. of polistine wasps (Hymenoptera: Vespi­ 144:63-150. dae). Z. Tierpsychoi. 68: 152-167. ---. 1991. The swarm-founding Polisti­ ---. 1985b. Colony development and so­ nae. Pages 191-231 in K. G. Ross and R. cial structure in a subtropical paper wasp, W. Matthews, eds. The social biology of Ropalidiafasciata (F.) (Hymenoptera: Ves­ wasps. Comstock Publication Associates, pidae). Res. Populo Ecoi. (Kyoto) 27: 333­ Cornell University Press, Ithaca, New 349. York. ---. 1986. Social behavior of Ropalidia KOnMA, J. 1984. Division oflabor and dom­ fasciata (Hymenoptera: Vespidae) females inance interaction among cofoundresses on satellite nests and on a nest with mul­ on pre-emergence colonies of Ropalidia tiple combs. J. Ethoi. 4: 73-80. fasciata (Hymenoptera: Vespidae). BioI. ---. 1987. Social behavior of the Aus­ Mag. (Okinawa) 22: 27-35. tralian paper wasp, Ropalidia revo/u­ MATSUURA, M., and SK. YAMANE. 1990. tiona/is (de Saussure) (Hymenoptera: Ves­ Comparative ethology of the Vespinae. pidae). J. Ethoi. 5: 115-124. Springer Verlag, Berlin. ---. 1993a. Behavior and social evolution Pardi, L. 1942. Recerche sui Polistini. 5. La of wasps: The communal aggregation hy- poliginia iniziale di Polistes gallicus (L.). 54 PACIFIC SCIENCE, Volume 49, January 1995

Boll. 1st. Entomol. Univ. Stud. Bologna nearly solitary paper wasp from Mada­ 14: 1-106. gascar (Hymenoptera: Vespidae). J. Kans. ---. 1946. Recerche sui Polistini. VII. La Entomol. Soc. 60: 549-556. "dominazione" e il ciclo annuale in Po­ ---. 1992. Extreme queen-worker di­ /istes gallicus (L.). Boll. 1st. Entomol. morphism in Ropa/idia ignobilis, a small Univ. Stud. Bologna 15: 25-84. colony wasp (Hymenoptera: Vespidae). SPRADBERY, J. P. 1991. Evolution of queen Insectes Soc. 39: 31-43. number and queen control in the Vespidae. WEST-EBERHARD, M. J. 1969. The social Pages 336-388 in K. G. Ross and R. W. biology of polistine wasps. Univ. Mich. Matthews, eds. The social biology of Mus. Zool. Misc. Publ. 140: 1-101. wasps. Comstock Publication Associates, ---. 1978. Polygyny and the evolution of Cornell University Press, Ithaca, New social behavior in wasps. J. Kans. Ento­ York. mol. Soc. 51: 832-856. SPRADBERY, J. P., and J. KonMA. 1989. Nest WILSON, E. O. 1971. The insect societies. descriptions and colony populations of Belknap Press of Harvard University eleven species of Ropa/idia (Hymenoptera: Press, Cambridge, Massachusetts. Vespidae) in New Guinea. Kontyu YAMANE, So. 1986. The colony cycle of the 57: 632-653. Sumatran paper wasp Ropa/idia (Icario/a) TURILLAZZI, S. 1980. Seasonal variations in variegata jacobsoni (Buysson), with refer­ the size and anatomy of Po/istes gallicus ence to the possible occurrence of serial (L.) (Hymenoptera: Vespidae). Monit. polygyny (Hymenoptera: Vespidae). Zool. Ital. (n.s.) 14: 63-75. Monit. Zool. Ital. (n.s.) 20: 135-161. TURILLAZZI, S., and M. TURILLAZZI. 1985. YAMANE, So., J. KonMA, and SK. YAMANE. Notes on the social behavior of Ropa/idia 1983. Queen worker size dimorphism in fasciata (F.) in West Java (Hymenoptera: an Oriental polistine wasp, Ropa/idia Vespidae). Monit. Zool. ltal. (n.s.) montana Carl (Hymenoptera: Vespidae). 19:219-230. Insectes Soc. 30: 416-422. WENZEL, J. W. 1987. Ropa/idia formosa, a

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