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Virgin queens in stingless bee (Apidae, Meliponinae) colonies: a review Vl Imperatriz-Fonseca, R Zucchi To cite this version: Vl Imperatriz-Fonseca, R Zucchi. Virgin queens in stingless bee (Apidae, Meliponinae) colonies: a review. Apidologie, Springer Verlag, 1995, 26 (3), pp.231-244. hal-00891261 HAL Id: hal-00891261 https://hal.archives-ouvertes.fr/hal-00891261 Submitted on 1 Jan 1995 HAL is a multi-disciplinary open access L’archive ouverte pluridisciplinaire HAL, est archive for the deposit and dissemination of sci- destinée au dépôt et à la diffusion de documents entific research documents, whether they are pub- scientifiques de niveau recherche, publiés ou non, lished or not. The documents may come from émanant des établissements d’enseignement et de teaching and research institutions in France or recherche français ou étrangers, des laboratoires abroad, or from public or private research centers. publics ou privés. Review article Virgin queens in stingless bee (Apidae, Meliponinae) colonies: a review VL Imperatriz-Fonseca R Zucchi 1 Departamento de Ecologia Geral, IBUSP, CP 11461, CEP 05422-970 São Paulo, SP; 2 Setor de Ecologia da Faculdade de Filosofia, Ciências e Letras da USP, CEP 14 300-000 Ribeirâo Preto, SP, Brazil (Received 9 December 1994; accepted 20 March 1995) Summary — There are no differences between Trigonini and Meliponini in terms of their treatment of virgin queens. They may stay in the nest, leave during swarm, supersede the dominant queen, or be killed, depending on what happens inside the colony. However, some kinds of behavior and strategies are characteristic of each species and part of its repertoire; examples are the onset of attractiveness, type of isolation, maturity, and the permanence of gynes in the colonies. Behavioral patterns are sim- ilar to all gynes. Attractive gynes have swollen abdomens, and abdominal glands are exposed during periods of attractiveness; they are very active, they run through the colony and search for trophal- laxis. Natural polygyny occurs in Melipona bicolor, but needs additional studies to be well understood. Temporary oligogyny occurs during supersedure process in Plebeia. The control of the number of vir- gin queens during certain periods of time is suggested in Trigona (Trigona), acting on the gynes’ emergence from royal cells or simultaneous metamorphosis of pupae. The presence of gynes in the nests stimulates swarm or supersedure. In these bees, one or more virgin queens depart with a swarm; fertilization occurs in a nuptial flight near the new nest. Swarm is a gradual process, with resource transportation from mother to daughter colonies. Supersedure may or may not be gradual. Attractive gynes, as well as workers, take active part in this process. Meliponinae / stingless bees / gyne / queen supersedure / swarm INTRODUCTION perenniality if the dominant queen dies. It is assumed that stingless bee queens have control over worker activities Meliponinae, which are highly eusocial bees, pheromonal and as occurs in other similar constitute a very interesting and diversified physiology, group. Little is known about their reproduc- eusocial species (Engels et al, 1987, 1993). tive strategies (Engels and Imperatriz-Fon- Although the nature of such regulations is seca, 1990). Massive provisioning of the unknown, it probably differs from the closely brood cells entails the need to keep gynes related Apinae. The permanence of attrac- (virgin queens) in the colony to ensure its tive gynes in the nests is a central point in stingless bees sociobiology. As stressed by BEHAVIOUR OF VIRGIN QUEENS Sakagami (1971), the socio-ethological IN MELIPONINI make-up of both societies (Apinae and that their similar Meliponinae) suggests The Meliponinae comprise 2 tribes: eusocial were reached patterns divergently. Meliponini, with a single genus, Melipona, The pattern of queen dominance pre- and Trigonini, with several genera (Moure, sented by Apis mellifera and its effect on 1961). The larger royal cells of the Trigo- workers is normally used for comparative nini distinguish them from the Meliponini, purposes. In this species, the queen con- whose queens, males and workers are bred trols the appearance of royal cells and also in identical cells. the activation of workers’ ovaries (Pain, In Meliponini, castes are determined by 1961; Free, In the 1987). Meliponinae, royal the interaction of genes with the environ- cells are constructed all the and year round, ment (Kerr et al, 1966; Velthuis and Som- whether the has influence on the queen any meijer, 1991). Virgin queens are smaller than construction of such cells is unknown. More- workers, they emerge all the year round, and over, the nurse-workers of most of the stud- are usually killed by workers. In Melipona, ied taxa have activated ovaries and pro- queens have similar characteristics in their duced that are eaten the eggs generally by behavioral repertoire; they have a shorter in such a if is queen. So, taxon, there any ontogeny than workers (data for M marginata control the it involves by queen, probably and M quadrifasciata), and normally emerge different in to pathways comparison Apinae. without attractiveness, ie they do not release This concerns the of work- mainly oviposition specific responses in workers. At the onset ers that will males, after dominant yield of attractivity they move more quickly around and the queen oviposition during opercula- the nest, requesting food from workers by tion of cells (Beig, 1972). making contact with their antennae aided by As regards the emerging virgin queens, their front legs. This is the stage at which they may be killed, supersede the dominant pheromone production probably starts. queen, or swarm to start a new nest with Attractive queens generally keep their workers. All 3 possibilities occur in different abdomen swollen and appear larger. They episodes of the colonial cycle. Several fac- actively seek the area where the new comb tors certainly influence these events, of is being built, particularly during the cell pro- which the respective promoters have not visioning and oviposition process, and may yet been identified. possibly ingest alimentary eggs laid by work- This paper presents a review of the virgin ers (Silva et al, 1972; Kleinert-Giovannini, queen’s behavior and permanence in sting- 1989). At the peak of their attractiveness, less bee nests. Most of the reproductive they expose their swollen abdomen to the of the aspects dealt with here are only factual and majority surrounding workers, through derive from unplanned events that took circular movements. Such gynes may then be killed or When the place in the course of several experiments accepted. gyne is and observations. This indicates the char- accepted, she helps workers eliminate the acter of this paper, which mainly intends to other virgin queens. report facts that are not normally delivered In normal colonies, the attractiveness of in formal publications on account of their the virgin queens causes an agonistic atti- incidental occurrence. The generic citations tude among workers who start chasing and throughout the text follow the classificatory killing them by twisting off their heads and system devised by Moure (1951, 1961) and other body parts. Such remains are placed Camargo and Pedro (1992). in the refuse area and removed from the colony. This is the fate of most Melipona M bicolor presents a natural polygyny. queens. M marginata, however, sometimes The term polygyny is used here in the same preserves a larger number of queens inside way as Herbers (1993) did: "to denote the the colony (Kleinert and Imperatriz-Fonseca, peaceful coexistence of 2 or more mated 1994). They take refuge in common areas, egg-layers (queens) in the same nest, ...and such as parts of the exoinvolucrum, remain- the intermixing of their offspring within one ing there for hours. From the body posture socially homogeneous unit". Colonies of M of the virgin queens, and the trophallaxis bicolorwith large populations may have as that occurs, it can be assumed that there is many as 5 physogastric queens simultane- an hierarchy among them. Therefore, in this ously involved in oviposition, apparently case, several queens are protected as a without any agonistic behavior (Bego, 1983, group by a physical barrier. 1988, 1989). There are even situations in Mota studied the development of abdom- which 1 queen remains to attend to the cell where the larval food is while inal glands in Melipona queens (Mota, 1982, being placed, 1988; Cruz-Landim and Mota, 1990). The another approaches and lays. Although presence and development of these glands dominance among queens is likely, conflicts may explain why the virgin queen rubs the are suggested by the male production by tip of her enlarged abdomen on the work- the workers, which is conspicuous than in ers’ head; this has been observed in polygynic nests, and the length of the inter- M quadrifasciata and M marginata at the val between oviposition by the queen and peak of her attractiveness. This might also cell operculation by the workers. Other explain the apparently defensive mecha- behaviors that differ are related to the kind nism used by the virgin queen of M quin- of courtship that workers pay to the queens, quefasciata, who spins around at high with pronounced antennal contact and a speed, hitting workers with her abdomen long duration. them into the distance. While and tossing Unlike those of other known stingless this is the mem- occurring, intersegmental bees, dominant queens of M bicolor rarely branes of the virgin queen’s abdomen are beat their wings, which thus do not wear out exposed, and whenever possible this region as they become older (Bego, 1988). A dom- is pressed against the workers’ heads (Zuc- inant queen may live as long as 7 years chi, 1977). Regarding the natural super- (Drumond, personal communication). Most sedure of queens in Melipona, Silva et al colonies kept in laboratories are mono- described the process for M quadri- (1972) Observations showed increased fasciata and M quinquefasciata.

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