Intrinsic colony conditions affect the provisioning and oviposition process in the stingless bee Melipona scutellaris

The cell provisioning and oviposition process (POP) is a unique characteristic of stingless bees (Meliponini), in which coordinated interactions between workers and queen regulate the filling of brood cells with larval resources and subsequent egg laying. Environmental conditions seem to regulate reproduction in stingless bees; however, little is known about how the amount of food affects quantitative sequences of the process. We examined intrinsic variables by comparing three colonies in distinct conditions (strong, intermediate and weak state). We predicted that some of these variables are correlated with temporal events of POP in Melipona scutellaris colonies. The results demonstrated that the strong colony had shorter periods of POP.

Pollen foraging in colonies of Melipona bicolor (Apidae, Meliponini): effects of season, colony size and queen number

We evaluated the ratio between the number of pollen foragers and the total number of bees entering colonies of Melipona bicolor, a facultative polygynous species of stingless bees. The variables considered in our analysis were: seasonality, colony size and the number of physogastric queens in each colony. The pollen forager ratios varied significantly between seasons; the ratio was higher in winter than in summer. However, colony size and number of queens per colony had no significant effect. We conclude that seasonal differences in pollen harvest are related to the production of sexuals and to the number of individuals and their body size.

The queen is dead-long live the workers: intraspecific parasitism by workers in the stingless bee Melipona scutellaris

Insect societies are well known for their high degree of cooperation, but their colonies can potentially be exploited by reproductive workers who lay unfertilized, male eggs, rather than work for the good of the colony. Recently, it has also been discovered that workers in bumblebees and Asian honeybees can succeed in entering and parasitizing unrelated colonies to produce their own male offspring. The aim of this study was to investigate whether such intraspecific worker parasitism might also occur in stingless bees, another group of highly social bees. Based on a large-scale genetic study of the species Melipona scutellaris, and the genotyping of nearly 600 males from 45 colonies, we show that similar to 20% of all males are workers` sons, but that around 80% of these had genotypes that were incompatible with them being the sons of workers of the resident queen. By tracking colonies over multiple generations, we show that these males were not produced by drifted workers, but rather by workers that were the offspring of a previous, superseded queen. This means that uniquely, workers reproductively parasitize the next-generation workforce. Our results are surprising given that most colonies were sampled many months after the previous queen had died and that workers normally only have a life expectancy of similar to 30 days. It also implies that reproductive workers greatly outlive all other workers. We explain our results in the context of kin selection theory, and the fact that it pays workers more from exploiting the colony if costs are carried by less related individuals.

The cuticular hydrocarbons profiles in the stingless bee Melipona marginata reflect task-related differences

Members of social insect colonies employ a large variety of chemical signals during their life. Of these, cuticular hydrocarbons are of primary importance for social insects since they allow for the recognition of conspecifics, nestmates and even members of different castes. The objectives of this study were (1) to characterize the variation of the chemical profiles among workers of the stingless bee Melipona marginata, and (2) to investigate the dependence of the chemical profiles on the age and on the behavior of the studied individuals. The results showed that cuticular hydrocarbon profiles of workers were composed of alkanes, alkenes and alkadienes that varied quantitatively and qualitatively according to function of workers in the colony. (C) 2010 Elsevier Ltd. All rights reserved.

Thoracic vibrations in stingless bees (Melipona seminigra): resonances of the thorax influence vibrations associated with flight but not those associated with sound production

Bees generate thoracic vibrations with their indirect flight muscles in various behavioural contexts. The main frequency component of non-flight vibrations, during which the wings are usually folded over the abdomen, is higher than that of thoracic vibrations that drive the wing movements for flight. So far, this has been concluded from an increase in natural frequency of the oscillating system in association with the wing adduction. In the present study, we measured the thoracic oscillations in stingless bees during stationary flight and during two types of non-flight behaviour, annoyance buzzing and forager communication, using laser vibrometry. As expected, the flight vibrations met all tested assumptions for resonant oscillations: slow build-up and decay of amplitude; increased frequency following reduction of the inertial load; and decreased frequency following an increase of the mass of the oscillating system. Resonances, however, do not play a significant role in the generation of non-flight vibrations. The strong decrease in main frequency at the end of the pulses indicates that these were driven at a frequency higher than the natural frequency of the system. Despite significant differences regarding the main frequency components and their oscillation amplitudes, the mechanism of generation is apparently similar in annoyance buzzing and forager vibrations. Both types of non-flight vibration induced oscillations of the wings and the legs in a similar way. Since these body parts transform thoracic oscillations into airborne sounds and substrate vibrations, annoyance buzzing can also be used to study mechanisms of signal generation and transmission potentially relevant in forager communication under controlled conditions.

The sound field generated by tethered stingless bees (Melipona scutellaris): inferences on its potential as a recruitment mechanism inside the hive

In stingless bees, recruitment of hive bees to food sources involves thoracic vibrations by foragers during trophallaxis. The temporal pattern of these vibrations correlates with the sugar concentration of the collected food. One possible pathway for transfering such information to nestmates is through airborne sound. In the present study, we investigated the transformation of thoracic vibrations into air particle velocity, sound pressure, and jet airflows in the stingless bee Melipona scutellaris. Whereas particle velocity and sound pressure were found all around and above vibrating individuals, there was no evidence for a jet airflow as with honey bees. The largest particle velocities were measured 5 mm above the wings (16.0 +/- 4.8 mm s(-1)). Around a vibrating individual, we found maximum particle velocities of 8.6 +/- 3.0 mm s(-1) (horizontal particle velocity) in front of the bee`s head and of 6.0 +/- 2.1 mm s(-1) (vertical particle velocity) behind its wings. Wing oscillations, which are mainly responsible for air particle movements in honey bees, significantly contributed to vertically oriented particle oscillations only close to the abdomen in M. scutellaris(distances <= 5 mm). Almost 80% of the hive bees attending trophallactic food transfers stayed within a range of 5 mm from the vibrating foragers. It remains to be shown, however, whether air particle velocity alone is strong enough to be detected by Johnston`s organ of the bee antenna. Taking the physiological properties of the honey bee`s Johnston`s organ as the reference, M. scutellaris hive bees are able to detect the forager vibrations through particle movements at distances of up to 2 cm.

Farnesoic acid O-methyl transferase (FAMeT) isoforms: Conserved traits and gene expression patterns related to caste differentiation in the stingless bee, Melipona scutellaris

Farnesoic acid O-methyl transferase (FAMeT) is the enzyme that catalyzes the formation of methyl farnesoate (MF) from farnesoic acid (FA) in the biosynthetic pathway of juvenile hormone (JH). This work reports the cloning, sequencing, and expression of FAMeT gene from the stingless bee Melipona scutellaris (MsFAMeT). The MsFAMeT in silica analysis showed that greatest sequence similarity is found in Apis mellifera and other insects, while relatively less similarity is shown in crustaceans. Evidence of alternative splicing of a 27 nucleotide (nt) microexon explains the presence of the detected isoforms, 1 and 2. The expression analysis of the two isoforms showed a marked difference when castes were compared, suggesting that they could be involved differently in the JH metabolism in M. scutellaris, providing new insights for the comprehension of female plasticity.

Espécies nectaríferas e poliníferas utilizadas pela abelha Melipona compressipes fasciculata (MELIPONINAE, APIDAE), no Maranhão

Este trabalho apresenta 79 espécies de plantas que fornecem néctar e/ou pólen para operárias de Melipona compressipes fasciculata, que é a abelha mais comum do Maranhão. A maioria dessas plantas são também visitadas por Apis mellifera. Sugere-se, com isso, o plantio de algumas espécies em estradas, cercas, ruas, avenidas e praças, a fim de melhorar a pasto apícola.

FONTES DE PÓLEN EXPLORADAS POR Melipona compressipes manaosensis(APIDAE: MELIPONINAE), ABELHA DA AMAZÔNIA CENTRAL

Analisou-se os pólen transportados por operárias de Melipona compressipes inanaosensisSchwarz, 1932, durante o período de um ano, entre agosto de 1988 ejulho de 1989. Foram encontrados 30 tipos polínicos distribuídos em 22 gêneros e 19 famílias. Das espécies vegetais coletadas pelas abelhas, as cássias representaram as principais fontes de pólen, seguidas de duas espécies de Miconiae de três espécies de Solarium.Outras espécies como Tapirira guianensis, Doliocarpussp., Lindackeriasp., Mimosa pudica,etc., foram bem coletadas pelas operárias, mas em períodos intercalados. Tudo indica que as coletas de pólen de Melipona compressipes inanaosensispodem sofrer influências com as mudanças climáticas, pois no período chuvoso, que correspondeu ao mês de maio, houve uma diminuição no número de espécies de plantas coletadas. Quando das chuvas, não foi observado operárias saindo ou retornando à colméia. As abelhas aproveitavam os intervalos de chuvas para retornarem as suas atividades externa.

Substituição natural de rainhas fisogástricas e distância de vôo dos machos em Tiuba (Melipona compressipes fasciculata Smith, 1854) e Uruçu (Melipona scutellaris Latreille, 1811) (Apidae, Meliponini)

Este trabalho relata detalhes da vida reprodutiva de duas espécies de abelhas sem ferrão. Rainhas velhas de Melipona compressipes fasciculata, no Maranhão, morrem e são substituídas com sucesso em todos os meses. Rainhas de Melipona scutellaris, trazidas de Lençóis (BA - nordeste do Brasil) para Uberlândia (MG, centro-sudeste do Brasil) morreram durante todos os meses e não mostraram a existência de trimestre preferencial para as novas rainhas iniciarem postura. Quarenta machos de M. scutellaris, após serem marcados no tórax e libertados em grupos de 10 a 100, 400, 800 e 1000 metros do meliponário, tiveram seus retornos observados. Todos os machos libertados a 100 e 400 metros regressaram ao meliponário, 7 de 10 machos e 2 de 10 machos retornaram de 800 e 1000 metros, respectivamente. Os machos esperam constantemente pela saída de uma rainha virgem, próximos às colônias órfãs, o que indica que a maioria das rainhas é inseminada próximo aos seus ninhos, portanto, a dispersão dos genes depende do vôo dos machos e da distância de enxameagem para ocupação de uma cavidade para o novo ninho.

Genetic breeding on the bee Melipona scutellaris (Apidae, Meliponinae)

A selection of queens of Melipona scutellaris through the most productive colonies were carried out during eight months in an orange honeyflow. Each of the colonies was evaluated by its production, that is, the gross weight production ( pollen, brood, geopropolis and wax of each hive). With this data a coefficient of repeatability was estimated by the intraclass correlation method, obtained r = 0.835 ± 0.071. The repeatibility is very high showing that the analysed data (production) is repeatable. Selection was then carried out using the regression coefficient of each colony and the respective production gain. Using these data the colonies were divided into three groups according to the method Vencovsky and Kerr (1982): a with the colonies of highest productivity, b of least productivity, and c of intermediary productivity. Colonies with the highest production (Group a) gave their queens to those of the lowest production (Group b) after their queens were taken out and killed; while those of intermediate (Group c) stayed with the same queens during the entire experiment both before and after the selection. The modifications in weight, that is, the genetic response was (R)= 7.98 gr per day which indicated a selection gain. The estimate of the realized herdability is twice the rate of the response to selection (R) by the selection differential (S2). That is then h²R=2(R/S2) then h²R= 0.166

Melitocoria de Zygia racemosa (Ducke) Barneby & Grimes por Melipona seminigra merrillae Cockerell, 1919 y Melipona compressipes manaosensis Schwarz, 1932 (Hymenoptera, Meliponina) en la Amazonía Central, Brasil

Durante el periodo de enero a abril de 2005 fueron encontradas semillas de Zygia racemosa, incorporadas al geoprópolis, en el interior de abejas sin aguijón en dos meliponarios experimentales en Manaus - AM. Fue observada intensa actividad de las obreras de M. seminigra merrillae y M. compressipes manaosensis llegando del campo con semillas fijas en las corbículas adheridas con resina y/o saliendo de las colonias con las semillas atrapadas en las mandíbulas. Ochenta semillas fueron recogidas en el interior de las colonias (mezcladas al geoprópolis en los basureros y fisuras) y también en el exterior, cerca de la entrada de las colonias, como resultado de la caída de esas semillas de las corbículas durante el vuelo de las obreras. Las semillas fueron plantadas en semilleros para la producción de esquejes y posterior identificación de la especie vegetal. Paralelamente se realizó el rastreo en campo en un radio de aproximadamente 3Km para la confirmación de la dispersión de las semillas, observaciones de comportamiento de las abejas forrajeando y recolección de semillas. Un total de 170 plántulas de Z. racemosa fueron encontradas en los alrededores del meliponario del GPA-INPA y 160 en el meliponario Vale Verde. Esos resultados indican que tanto M. compressipes manaosensis como M. seminigra merrillae recolectan y dispersan las semillas de Z. racemosa.

Five egg-laying queens in a single colony of brazilian stingless bees (Melipona scutellaris Latreille

Polygyny, characterized by the presence of several egg-laying queens, is considered as a temporary colony status. In stingless bees it is rarely observed. This paper reports the first case of natural polygyny in Melipona scutellaris colony, with five egg-laying queens.

Espectro polínico de mel de tiúba (Melipona fasciculata Smith, 1854, Hymenoptera, Apidae)

O espectro polínico do mel de Melipona fasciculata Smith foi analisado com o objetivo de identificar os recursos nectaríferos utilizados por essa espécie. A identificação das plantas visitadas foi realizada com base na análise dos tipos polínicos encontrados em 12 amostras de mel coletadas, mensalmente, em uma colônia localizada no município de Palmeirândia, na área da Baixada Ocidental Maranhense (02º40'47,6S, 44º52'39,8"W), Brasil. As análises quantitativas e qualitativas foram realizadas com o objetivo de determinar as porcentagens e classes de frequência dos tipos polínicos presentes nas amostras de mel. Foram encontrados 45 tipos polínicos, sendo Pontederia parviflora Alexander (Pontederiaceae), espécie mais frequente em todo o período de amostragem (38,6%), pólen dominante em Outubro (86%), Junho (85%), Julho (76%), Agosto (49%) e Setembro (51%) e pólen acessório em Dezembro, Janeiro e Março. Mimosa caesalpiniifolia Benth (Mimosaceae) foi a segunda espécie mais frequente (22,8%) sendo pólen dominante em Novembro (46%), Abril (74%) e Maio (72%). Myrcia eximia DC (Myrtaceae) foi considerada pólen isolado importante. As famílias mais representativas no espectro polínico das amostras de mel foram Pontederiaceae e Mimosaceae. 50% dos méis foram biflorais, havendo também méis monoflorais (41,7%) e heteroflorais (8,3%).

Formação das castas no gênero Melípona (Illiger, 1806)

The present work is destinated to prove that the castes : workers and queens, in Melipona bees are due to genetic factors and not to differences in food. 2) Material used: Hives of Melipona quadri-fasciata anthidioides (Lep. 1836), M. schenki schenki (Gribodo, 1893), M. fasciata rufiventris (Lep. 1836), M. quadri-fasciata vicina (Lep. 1836), M. marginata marginata (Lep. 1836), Apis mellifera (L. 1758). 3) It should be pointed out that in Melipona bees there are no royal cells for the queens, but all the cells are of the same size independently of being destinated for workers, queens or drones. The numerous queens which are born are killed soon after emerging from their cells. 4) Changes of feeding in quality and in quantity caused no variation of castes. The only variable factor is the size, which becomes bigger when the bee is well nourished. 5) The offsprings of 5 hives were examined : 3 of M. quadri-fasciata anthidioides (n.o 1, n.o 2 and n.o 3), 1 of M. quadri-fasciata vicina (n.o 4) and 1 of M. marginata marginata (n.o 5). Combs of about 40 cells were taken into laboratory and the type of bee registered immediately after emerging. The results of the counts were: BOX COMB WORKER QUEEN PERCENTAGE Σ X2 to 12,5% Nº 1 1th 69 8 10,4% 0, 3139 " 1 2nd 144 18 11,1% 0, 2856 " 2 1th 52 8 13,3% 0, 0384 " 3 1th 45 10 18,2% 1, 6736 " 4 1th 56 4 6,7% 1, 8686 " 4 2nd 29 4 12,1% 0,00432 Σ X2 to 25% " 5 1th 34 14 29,2% 0,44444 "5 2nd 83 27 24,5% 0, 0121 In the 4 first boxes there is a percentage of 11,63% queens and in the last there is a percentage of 25,95%. 6) These percentages are very near two genetical ratios: 12,5% or 7:1, and 25% or 3:1, which correspond to a trifactorial and a bifactorial back-cross. Carrying out a X² test no significant deviations were found ( X² to 12,5% and to 25% and table 1 to 4). 7) We suppose that the formula for the queen in the first case (11,65%) is: AaBbCc. Since the Melipona bees are arrhenotokous hymenopteres, the drones are haploid and may have any one of the following eight formulas, corresponding to the gonic segregation of the queem : ABC, ABc, Abc, Abc, AbC, aBC, aBc, abC, abc. Anyone combination of these males with the queen will give a segregation of 7 workers to 1 queen, since there is always only one triple heterozygote among the eight possible segregates (table 5). 8) In order to explain the second case, it is suffient to assume that in this species there are only two pairs of factors, the queen being the double heterozygote : AaBb, while the drones may have any one of the following constitutions: AB, Ab, aB and ab. Workers are again all diploids which are homozygous for one or both factors, for instance: AABB, AABb, AaBB, aaBb, AAbb, etc. (table 6). 9) It is suggested that the genus Melipona is an intermediary type between the solitary bees, where all females are fertile independently of their feeding, and the genera Apis and Trigona, where without special feeding all females are born sterile, while only specially fed females develop into fertile queens. 10) No speculations are put forward with regards to the evolutionary mechanism which may have been responsible for the development of the genetical determination of castes in Melipona, since it seems advisable point to extend the studies to other insects with complicated caste systems.