1996

WILMS, W.; IMPERATRIZ-FONSECA, V.L. & ENGELS, W.1996. Resource partitioning between highly eusocial bees and possible impact of the introduced honeybee on native stingless bees in the Brazilian Atlantic rainforest. Studies on Neotropical Fauna and Environment 31 (3-4): 137-151.

Abstract:The highly eusocial bee community of the neotropical Atlantic Rainforest was studied at Boraceia Biological Station in the state of Sao Paulo, Brazil. In this reserve, 17 species of stingless bees and the introduced Africanized honey bee were found, the latter being the most abundant flower visitor. Of all flowering plants, Asteraceae and Myrtaceae were particularly important as resources for bees. Trophic niche overlap between the various species of stingless bees is evident and it was generally larger within the tribes Meliponini and Trigonini than between members of different tribes. Nevertheless, in the stingless bee community the competitive pressure is rather uniformly spread. The trophic niche of the Africanized honey bee can be positioned between those of Meliponini and Trigonini. Today this introduced species represents the main competitor in this bee community. However, its impact on native stingless bee populations is apparently buffered by mass-flowering trees which are the most important food plants of the indigenous highly eusocial bees.

Keywords:Atlantic Rainforest of Brazil; bee community; stingless bees; Africanized honey bee; nutritional resource partitioning; mass-flowering

1997

IMPERATRIZ-FONSECA, V.L.; CRUZ-LANDIM, C. & MORAES, R.L.M.S. 1997. Dwarf gynes in Nannotrigonatestasceicornis (Meliponinae). Behaviour, exocrine gland morphology and reproductive status. Apidologie28 (3-4):113-122.

Abstract:The behaviour and morphology of dwarf gynes produced in worker-sized cells of nor mal colonies in Nannotrigonatestaceicornis (Meliponinae, Trigonini) were studied. The behaviour of these dwarf virgin queens was the same as observed for normal Trigonine gynes. The glandular equipment is also the same: Dufour glands, fat bodies and spermathecae are present. Despite these similarities, their ovaries are different. The functional significance of dwarf gynes is unknown, but may be a basis for an alternative reproductive strategy.

Keywords:dwarf gynes; larval food; reproductive strategies; stingless bees; Trigonini; Nannotrigonatestaceicornis

VIANA, B.F.; KLEINERT, A.M.P.IMPERATRIZ-FONSECA, V.L. 1997.Abundance and flower visits of bees in a cerrado of Bahia, tropical Brazil. Studies on Neotropical Fauna and Environment 32(4):212-219.

Abstract:Abundance, seasonal phenology and flower visits of eusocial bees were studied in Diamantina, a national park with cerrado vegetation in Bahia, tropical Brazil. About 700 bees were collected, mainly native stingless bees and the introduced Africanized honey bee. Sampling along a transect was most effective March through September during the dry season. The foraging worker bees were observed on flowers of over 60 angiosperm species of which a few were visited with high frequency. Foraging activity concentrated on flowering plants of the families Leguminosae and Asteraceae. The results are discussed under aspects of nutritional resource partitioning by bee communities in neotropical habitats and the specific composition of a cerrado apifauna.

Keywords:eusocial bee community; Apidae; resource partitioning; cerrado vegetation; flower visits; seasonal abundance; Bahia; Brazil

1998

ALONSO, W.J.; LUCENA, T.F.; FRACASSO, C.M.; VELTHUIS, H.H.W. & IMPERATRIZ-FONSECA, V.L.1998. Do Meliponabicolor workers distinguish relatedness among different physogastric queens? Apidologie29(6):503-512.

Abstract:The prediction that the polygynous stingless bee Melipona bicolor workers should distinguish physogastric queens to whom they have different degrees of relatedness was tested. One colony with two physogastric queens was monitored for 5 consecutive days for the occurrence of the provision and oviposition processes (POP) of individually marked workers. We recorded cell provisioning, egg laying, and cell sealing. The presence of one or both of the queens during each POP was recorded. After that, one of the queens from the colony used in the first experiment was removed from the colony and substituted by two other non-related physogastric queens, from other colonies. In both cases preferences by workers towards their mother or aunt or to the unrelated queens were not detected. In a third experiment we demonstrated that guard bees at entrance of a colony do not reject a physogastric foreign queen. Evolutionary and husbandry implications are discussed.

Keywords:kin recognition; polygyny; provision and oviposition processes; Melipona bicolor

CRUZ-LANDIM, C.; REGINATO, R.D. & IMPERATRIZ-FONSECA, V.L.1998. Variation on ovariole number in Meliponinae (Hymenoptera, Apidae) queen’s ovaries with comments on ovary development and caste differentiation. Papéis Av. de Zool.40(18): 289-296.

Abstract:The present paper concerns the variation of the number of ovariole per ovary in the females of some Meliponinae species. In workers, the number of ovarioles was always of 4 per ovary, but in some species the queens had a variable number of ovarioles, from 4 to 15 per ovary (as in Plebeia remota, Nannotrigona testaceicornis, Trigona spinipes, Schwarziana quadripunctata). The variation of the number of ovarioles was also found in the same species from individual to individual and among the ovaries of a single individual.

Keywords:ovariole number; stingless bees; queen; worker

IMPERATRIZ-FONSECA, V.L.KLEINERT, A.M.P.1998. Worker reproduction in the stingless bee Friesellaschrottkyi (Apidae, Meliponinae). Entomologia Generalis 23(3):169-175.

Abstract:In Friesella (Trigona) schrottkyi Moure 1946, workers do not lay alimentary eggs in the presence of the queen, but functional reproductive eggs are laid in short periods of their adult life. The here reported observations focus on one of these periods, that began just after an accidental gyne's escape from ist chamber (in a colony maintained inside the Bee Lab), and was divided in 2 phases, before (17 days) and after (16 days) swarm departure. Worker ovipositions occurred during the sealing of the cell or by worker manipulation of closed cells. The discussion of this phenomenon is centered on queen control over laying of reproductive eggs by workers.

Keywords:Friesella (Trigona) schrottkyi Moure 1946; pattern of oviposition; virgin queens; swarming; supersedure

IMPERATRIZ-FONSECA, V.L.; MATOS, E.T.; NOGUEIRA-FERREIRA, F.H. & VELTHUIS, H.H.W. 1998. A case of multiple mating in stingless bees. Insectes Sociaux45(2):231-233.

Abstract:In several stingless bee species many males aggregate in the vicinity of a nest when a virgin queen is present in the colony and is preparing for the nuptial flight. We report such male assemblage for Tetragonisca angustula. The departure of a virgin queen from the colony and the subsequent mating could be video-recorded, because the queen and the males that had mounted her fell to the ground. Since at least two males had lost their genitalia, multiple mating seems to have occurred. This is in contrast with the prevailing view found in literature concerning the mating biology of stingless bees.

Keywords:multiple mating; stingless bees; Tetragonisca angustula; Meliponinae

PETERS, J.M.; QUELLER, D.C.; IMPERATRIZ-FONSECA, V.L. & STRASSMANN, J.E. 1998. Microsatellite loci from the stingless bees. Molecular Ecology7:783-92.

RAMALHO, M.; IMPERATRIZ-FONSECA, V.L. & GIANNINI,T.C. 1998. Within-colony size variation of foragers and pollen load capacity in the stingless bee Meliponaquadrifasciataanthidioides(Apidae, Hymenoptera). Apidologie29(3):221-228.

Abstract:Within-nest worker size variation in the eusocial stingless bees is a contingent phe nomenon of unknown adaptive value. We assume that the magnitude of variation represents a compromise between a minimum population of foragers and foraging efficiency at colony level. In Melipona quadrifasciata, worker size was found to vary according to colony conditions, and the pollen carrying efficiency of an individual was related to its size. On average, the foragers from a weak colony are smaller and are able to carry greater amounts of pollen per unity of body weight ('load capacity') than the larger foragers from a strong colony. The allometric variation of the corbicula (the pollen carrying structure in the hind tibia) contributes to the observed decrease in pollen load capacity with increased body size. By higher pollen intake per worker, the colonies with smaller bees could increasing the rate of brood production and colony population recovery after population crashes.

Keywords:Melipona quadrifasciata; stingless bees; worker size; pollen load; allometric variations

1999

KOEDAM D; CONTRERA, F.A.L.; & IMPERATRIZ-FONSECA, V.L.1999. Clustered male production by workers in the stingless bee Meliponasubnitida. Insectes Sociaux46(4):387-391.

Abstract: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.

Keywords:stingless bees; Melipona subnitida; worker reproduction; male production; queen dominance

PETERS, J.M.; QUELLER, D.C.; IMPERATRIZ-FONSECA, V.L.; ROUBIK, D.W. & STRASSMANN, J.E. 1999. Mate number, kin selection, and social conflicts in stingless bees and honey bees. Proceedings of the Royal Society of London Series B-Biological Sciences 266(1417):379-384.

Abstract:Microsatellite genotyping of workers from 13 species (ten genera) of stingless bees shows that genetic relatedness is very high. Workers are usually daughters of a single, singly mated queen. This observation, coupled with the multiple mating of honeybee queens, permits kin selection theory to account for many differences in the social biology of the two taxa. First, in contrast to honeybees, where workers are predicted to and do police each other's male production, stingless bee workers are predicted to compete directly with the queen for rights to produce males. This leads to behavioural and reproductive conflict during oviposition. Second, the riskthat a daughter queen will attack the mother queen is higher in honeybees, as is the cost of such an attack to workers. This explains why stingless bees commonly have virgin queens in the nest, but honeybees do not. It also explains why in honeybees the mother queen leaves to found a new nest, while in stingless bees it is the daughter queen who leaves.

Keywords:kin selection; mate number; social conflict; microsatellite; Apis; Meliponini

VELTHUIS, H.H.W.; KOEDAM, D. & IMPERATRIZ-FONSECA, V.L.1999. The rate of brood cell production in the stingless bee Meliponabicolor fluctuates with nest box temperature. Revista de Etologia2(1):141-145.

Abstract:The nesting sites of stingless bees probably offer the possibility of maintaining quite a stable temperature for the developing brood. For laboratory studies, the nests of stingless bees are often placed in observation hives, which are made of wood and have glass lid on top. In this paper, we report on the effect of glass plate removal on brood cell production in Melipona bicolor and demonstrate the impact of temperature fluctuations on the brood cell activity of this species. The production of new brood cells was observed in a polygynous colony which was experimentally subject to different regimes of cold and warm episodes. These results were compared with data obtained simultaneously form three other colonies, each in this own incubator box. It showed that the rate of brood cell production during cold episodes was only 50-60% that during the warmer ones. The observations lead to several recommendations form improving conditions under which colonies of stingless bees are kept and studied.

Keywords:thermoregulation; brood cell production; stingless bees;Melipona bicolor

2000

HILÁRIO, S.D.; IMPERATRIZ-FONSECA, V.L.; KLEINERT, A.M.P.2000. Flight Activity and colony strength in the stingless bee Meliponabicolorbicolor (Apidae, Meliponinae). Revista Brasileira de Biologia60(2):299-306.

Abstract:Flight activity of Melipona bicolor bicolor, coming from Cunha (23o05'S, 44o55'W), AtlanticForest, was studied in ten colonies, and in two periods: from July to September 1993 and from August to September 1995. The colonies were grouped in weak, medium and strong, according to the diameter of the combs, which can provide a good idea of the number of cells built. 855 observations were accomplished for 5 minutes, every half-hour, from 8 to 18 hours. The total number of bees that entered and left the hive and the number of bees that arrived with mud, pollen and resin, besides the number that went out with debris in that period were counted. It was also registered the temperature and the relative humidity of the air. The total external activity, as well as pollen collection, was maximum in the first hours of the morning, mainly in strong colonies. Weak colonies moved their maximum activity approximately to 12 hours. Pollen collection declined gradually, while mud and resin collection rose; removal of debris was greater in the beginning of the morning and in the end of the afternoon. Flight activity increased as relative humidity of the air rose, being optimum for strong colonies in the range between 80%-89%, and for the weakest colonies between 70%-79%. The minimum temperature observed for exit of the bees was 11oC, with optimum temperatures ranging between 17oC and 22oC. The results showed that the general state of the colony influences the different strategies of food collection and that these bees should be adapted to environments of high relative humidity as the Atlantic forest.

Keywords: stingless bees; flight activity; climate conditions; resource collection

FRANCKE, W.; LUBKE, G.; SCHRODER, W.; RECKZIEGEL, A.; IMPERATRIZ-FONSECA, V.L.; KLEINERT, A.M.P.; ENGELS, E.; HARTFELDER, K.; RADTKE, R. & ENGELS, W. 2000. Identification of oxygen containing volatiles in cephalic secretions of workers of Brazilian chemical society. J. Brazilian Chemical Society11(6):562-571.

Abstract: The volatile constituents of cephalic secrections of 11 Brazilian social stingless bee species of the Tetragonisca - Tetragona line have been analysed. By gas chromatography/mass spectrometry 145 compounds could be identified which include 72 esters, 22 alcohols, 16 carboxylic acids, 13 terpenoids, 8 aldehydes, 7 ketones, 4 aromatic compounds, 2 lactones and 1 dihydropyran. Structural relations, origin, and distribution of these compounds are discussed. With respect to qualitative and quantitative composition, each species shows a specific odour pattern which is made up by less specific components. To a certain extent, closely related species show some similarities in the odour bouquets. The mass spectrometric fragmentation patterns of typical wax type esters and DMDS derivatives of unsaturated esters are discussed in detail.

Keywords: stingless bees; cephalic secretions; Tetragonisca; Tetragona.

2001

CRUZ-LÓPEZ, L.; PATRÍCIO, E.F.L.R.A.; & MORGAN, E.D. 2001. Secretions of stingless bees: the Dufour Gland of Nannotrigona testaceicornis. Journal of Chemical Ecology 27(1): 69-80.

Abstract: The Dufour gland of Nannotrigona testaceicornis is a large, wide, pear-shaped sac. The gland secretion consists chiefly of the diterpene ester all-trans-geranylgeranyl acetate (64% of the total), together with a complex mixture of small amounts of cyclic ketals; mono-, sesqui-, and diterpene compounds; acetates; and other oxygenated compounds. Samples of N. testaceicornis collected at two sites in Brazil and one in México shared the same composition of their glands, suggesting that the species is uniform over this wide geographical area.

Keywords: Hymenoptera;Apidae;Meliponinae;exocrine secretion;geranylgeranyl acetate;terpene;oxidation products

HILÁRIO, S.D.; IMPERATRIZ-FONSECA, V.L.KLEINERT, A.M.P.2001. Responses to climatic factors by foragers of Plebeia pugnax Moure (in litt.) (Apidae, Meliponinae). Revista Brasileira de Biologia61(2):191-196.

Abstract: Flight activity of Plebeiapugnax Moure (in litt.) was studied in six colonies coming from Cunha, SP, from July to October 1994. Twice a week, from 8:00 a.m. to 6:00 p.m., for 5 minutes every half-hour, all the bees entering and leaving the hives were counted. Six hundred counts were made and the materials that foragers carried were recorded. Data were analysed in relation to temperature, relative humidity, light intensity and day time. Foragers' flight activity was relatively constant in a wide range of temperature, from 22°C to 34°C. The minimum temperature for the beginning of flight activity was 14°C. Effective flight activity (when foragers of all colonies were leaving the hives) occurred at 15°C. These bees also flew within a wide range of relative humidity, from 30% to 100%, decreasing slowly after 50%. Flight activity increased as light intensity rose and it has also increased as the hours passed by, reaching a peak around midday and decreasing gradually afterwards. Pollen was collected all day long, while resin collection was relatively constant and debris transportation was slightly higher after 10:00 h. From all known Plebeia species, this one flew on the lowest temperature ever registered for this genus.

Keywords: flight activity; stingless bees; climatic factors

VELTHUIS, H.H.W.; ROELING, A. & IMPERATRIZ-FONSECA, V.L.2001.Repartition of reproduction among queens in the polygynous stingless bee Meliponabicolor. Proc. Exper. Appl. Entomol. NEVAmsterdam12:45-49.

Abstract: The mechanisms leading to reproductive skew among queens in a polygynous colony of Melipona bicolor were studied. Queens differ in the probability of remaining the sole queen at a cell during the provisioning and oviposition process. Being alone enhances the probability of receiving a trophic egg. When together with another queen egg-laying is random. Therefore, reproductive dominance is the consequence of differential food uptake, not of a dominance hierarchy.

Keywords: polygyny; reproductive skew; trophic eggs;Melipona bicolor

KOEDAM, D.; VELTHUIS, H.H.W.; DOHMEN, M.R. & IMPERATRIZ-FONSECA, V.L.2001. The behaviour of laying workers and the morphology and viability of their eggs in Meliponabicolorbicolor.Physiological Entomology 26 (3): 254-259.