-Female behaviour in social Hymenoptera and the queen-worker conflict with respect to male production have been the focus of many studies. Although male production is an investment that is in conflict with investment in colony size, males play a vital role in colony reproduction. This paper reviews the production patterns of male stingless bees, their activities once they have reached adulthood and their origin (i.e., are they sons of workers or of queens). The existence of a broad spectrum of species-specific patterns of male production, sex ratios, and male parentage offers ample opportunities to discuss the influence of ecology on the dynamics of stingless bee colony life. The paper also argues that selfishness causes the queen and the workers to compete and each to adopt certain strategies in their effort to produce male progeny. It is this competition, expressed in various forms during the characteristic and socially complex process of cell provisioning and oviposition, that could help explain the variable outcomes of male parentage at the species level as we currently know them. stingless bee male / sex ratio / life history / male aggregation / queen-worker conflict / provisioning and oviposition process

In stingless bees brood cells are sequentially filled with liquid larval food (mass-provisioning), upon which the queen lays an egg. Thereafter the cell is closed by a worker. This study showed that during these processes workers of Melipona subnitida regularly laid eggs that served as food for the queen. Occasionally cells were oviposited in and immediately closed by a worker. These cells always rendered males. Some of these reproductive workers were seen to lay a trophic egg as well. Cells which were exclusively oviposited in by the physogastric queen gave rise to workers and queens only. In one colony it could be verified that three workers alone, which differed in age by one day, laid 15 male-producing eggs within a period of two successive weeks. Among them the number of ovipositions was positively related to the order in which workers eclosed -the oldest worker laying most eggs -and inversely related to the number of times they closed cells oviposited in exclusively by the queen. Apparently the physogastric queen was not able to stop certain workers from reproducing. We therefore conclude that some workers in M. subnitida temporarily dominated their queens in egg-laying.

Abstract. To study the reproductive potential of workers in the stingless bee Trigona (Tetragonisca) angustula Illiger (Apidae, Meliponinae), we examined the morphological quality of their eggs. Worker‐eggs were all placed on the inside of the upper wall of brood cells. Normally, such eggs are consumed by the queen and are therefore referred to as trophic. The provisioning and oviposition processes in queenright colonies are characterized by the occurrence of circular aggregations of workers, ‘rosettes’, around the broodcell opening immediately after the release of a worker‐egg. The ovaries of 35% of these rosette workers contained mature, chorionated eggs. In a single worker ovary, always only one mature egg was found. Some of the eggs, dissected from the ovarioles, showed a reticulate chorion pattern. Worker‐eggs which lacked this pattern were significantly bigger than patterned eggs. Scanning electron micrographs revealed that the patterned worker‐eggs are similar in appearance to queen‐eggs. After a worker‐egg had been removed experimentally from a broodcell, the same cell could be oviposited by a worker again. Light microscopic analysis revealed that all these worker‐eggs lacked the reticulate chorion pattern and were very similar in their morphology. In a colony without a laying queen workers laid eggs which had a reticulate pattern on the chorion. Since these eggs developed into males, we assume that the pattern on the chorion is characteristic for reproductive eggs. We also assume that the queen prevents the release of reproductive eggs by the workers. However, she does not inhibit the development of this type of egg.

Several recent hypotheses, including sensory drive and sensory exploitation, suggest that receiver biases may drive selection of biological signals in the context of sexual selection. Here we suggest that a similar mechanism may have led to convergence of patterns in flowers, stingless bee nest entrances, and pitchers of insectivorous plants. A survey of these non-related visual stimuli shows that they share features such as stripes, dark centre, and peripheral dots. Next, we experimentally show that in stingless bees the close-up approach to a flower is guided by dark centre preference. Moreover, in the approach towards their nest entrance, they have a spontaneous preference for entrance patterns containing a dark centre and disrupted ornamentation. Together with existing empirical evidence on the honeybee's and other insects' orientation to flowers, this suggests that the signal receivers of the natural patterns we examined, mainly Hymenoptera, have spontaneous preferences for radiating stripes, dark centres, and peripheral dots. These receiver biases may have evolved in other behavioural contexts in the ancestors of Hymenoptera, but our findings suggest that they have triggered the convergent evolution of visual stimuli in floral guides, stingless bee nest entrances, and insectivorous pitchers.

No abstract

Several insect taxa have evolved symbioses with actinobacteria that protect the host or its nutritional resources against pathogens. Digger wasps of the genus Philanthus (‘beewolves’; Hymenoptera, Crabronidae) cultivate ‘Candidatus Streptomyces philanthi’ (Ca. S. philanthi) in specialized antennal glands and transfer them to subterranean brood cells, where the symbionts provide protection for the beewolf larva against pathogens by producing a cocktail of antibiotic substances. The present study investigates the occurrence of antennal symbionts in two species of the genus Trachypus, which is the closest relative to Philanthus. Cross sections of the antennae from females of both Trachypus denticollis and Trachypus boharti reveal bacteria‐containing gland reservoirs that are morphologically very similar to those found in Philanthus spp. Polymerase chain reaction‐based screens with specific primers, sequencing of the partial 16S rRNA gene, and fluorescence in situ hybridization confirm the presence of close relatives of ‘Ca. S. philanthi’ in T. denticollis and T. boharti. However, 16S rRNA sequence divergence between Trachypus spp. and Philanthus spp. symbionts is much higher for T. boharti than for T. denticollis, suggesting that horizontal transmission and/or de novo uptake of symbionts from the environment occurs occasionally. The results obtained indicate that the protective symbiosis with antennal actinobacteria is older and more widespread than previously recognized and occurs in at least two genera of digger wasps that comprise approximately 170 species.

Abstract. As in many other stingless bees, Melipona bicolor bicolor Lepeletier (Apidae: Meliponinae) workers lay two morphologically distinct types of eggs: slender ones that have a typical patterned chorion, and larger ones that lack this pattern. In this paper we report on the relation between egg morphology and the behaviour of the workers that lay such eggs. In most cases, the laying of each of these egg types is accompanied by a unique sequence of behaviours. After a worker has laid the unpatterned type of egg, she generally leaves the cell, giving the queen the possibility of eating this egg. In the case of the patterned egg type, the worker usually closes the cell immediately after her egg laying. When worker egg laying occurs right after a series of regurgitations, it stops the queen from ovipositing. When, instead, a worker lays an egg after queen oviposition, the cell contains two eggs. This study also revealed cases in which workers laid slender, patterned eggs without closing the cell, and other cases where workers laid large, spherical, unpatterned eggs and instantly closed the cell. Experiments in which worker eggs, destined to be eaten by the queen, were protected by covering the cell artificially with a piece of wax showed that some of these eggs developed into larvae, although they were occasionally relatively small. The occurrence of a range of combinations of egg‐laying behaviours and egg morphologies in M. b. bicolor workers is discussed from the perspective of worker egg laying in other stingless bees.

Potential conflict between the queen and workers over the production of males is expected in stingless bees as a result of the higher relatedness of workers with their sons than with their brothers. This conflict was studied in Melipona subnitida by observing how the queen and the workers share in male production. The oviposition of individual cells was observed in two colonies with individually marked workers for a period of 51 and 40 days respectively. The gender that developed from these cells was then determined. The results revealed that most male production was concentrated in a 2 -3-week period, during which laying workers were present. During these weeks, the queens produced twice as many males as all laying workers together. Outside this distinct period, the queens produced an occasional male. A reproductive worker either oviposited before the queen did, in which case she immediately proceeded to close the cell and thus prevented the queen from oviposition, or oviposited and sealed the cell after the queen had laid an egg. When cell construction and oviposition occured on several combs simultaneously, the workers preferentially laid male eggs on the newest combs. We discuss the proximate mechanism and ultimate cause of the way in which queenworker male production occurred. In conclusion, we argue that overt behavioural conflict, occasionally displayed by reproductive workers of this species, can be of great cost to the colony.