Juvenile hormone affects age polyethism, ovarian status and cuticular hydrocarbon profile in workers of the wasp Polybia occidentalis.

Division of labor is one of the most striking features in the evolution of eusociality. Juvenile hormone (JH) mediates reproductive status and aggression among nestmates in primitively eusocial Hymenoptera (species without morphologically distinct castes). In highly social species it has apparently lost its gonadotropic role and primarily regulates the division of labor in the worker caste. Polybia occidentalis, a Neotropical swarm-founding wasp, is an ideal model to understand how JH levels mirror social context and reproductive opportunities because of the absence of a clear morphological caste dimorphism. In this study, we tested the hypothesis that JH influences division of labor, ovary activation and cuticular hydrocarbon profiles of workers. Our observations confirmed that a JH analog (methoprene) and an inhibitor of JH biosynthesis (precocene) affected the cuticular chemical profile associated with age polyethism. Also, methoprene and precocene-I treatment of females influenced ovarian activation differently (individuals treated with methoprene expressed more activated ovaries while precocene treatment did not have significant effect). These results suggest that different hormonal levels induce a differential expression of cuticular chemicals associated with workers' age polyethism, which may be essential for keeping the social cohesion among workers throughout their lives in the colony. Furthermore, JH is likely to play a gonadotropic role in P. occidentalis. JH has apparently undergone certain modifications in social Hymenoptera, presenting multifaceted functions in different species.

Osmophores of Caryocar brasiliense (Caryocaraceae): a particular structure of the androecium that releases an unusual scent.

Caryocar brasiliense is a flagship species of the Brazilian Cerrado. It produces flowers with a strong peculiar scent, which are pollinated by bats and occasionally moths with short mouthparts. However, the cues responsible for attracting these nocturnal pollinators remain unknown. We aimed to identify osmophores of C. brasiliense, describe the ultrastructure of the cells involved in the synthesis and release of floral odour, and identify the constituents of the floral bouquet. We performed field observations and histochemical and ultrastructural analyses of flowers focusing on the androecium. Gas chromatography-mass spectrometry was used to analyse the scents emitted. Filament epidermal cells were found to possess an unusual shape and be responsible for the main production and release of odour. These cells, called foraminous cells, are elongate and possess pores where their cell walls are abruptly thin. The cuticle is practically absent over the pores, which facilitates odour emission. The foraminous cells have conspicuous nuclei and organelle-rich cytoplasm where oil droplets can be seen prior to anthesis. The features of these cells remain similar during anthesis, but many vesicles fuse with the plasma membrane and the number of oil droplets in the cytosol decreases. Twenty-nine components were found in the scent, especially fatty acid derivatives and N- and S-bearing compounds. Our analyses revealed that the androecium of C. brasiliense has a particular structure that acts as an osmophore. The scent from the androecium resembles that of the entire flower, which is an unprecedented finding for a plant with single flowers as the pollination unit.

Scientific Note: Warming Nurses, a New Worker Role Recorded for the First Time in Stingless Bees.

Nest temperature is a crucial variable that determines colony survival in social insects. The successful production and development of a new brood, therefore, depends on stable thermal conditions and limited temperature oscillations. Thermoregulatory processes are useful in controlling both individual activity and for the maintenance of colony temperature. We measured heat production generated by nurse bees working on brood combs of the stingless bee Melipona scutellaris (Hymenoptera, Apidae, Meliponini) in this study; our results enabled us to identify the existence of a new task performed by nurse bees, referred to here as 'hot bees' because of their higher thorax surface temperature (4°C above that of other bees within the brood area). This additional heat has been little studied in stingless bees but is likely the result of thorax muscle contractions or, indeed, the development of this musculature as these are recently emerged individuals. We hypothesize that these 'hot bees' contribute to the maintenance of warmth within the nest brood area.

Queen signals in a stingless bee: suppression of worker ovary activation and spatial distribution of active compounds.

In most species of social insect the queen signals her presence to her workers via pheromones. Worker responses to queen pheromones include retinue formation around the queen, inhibition of queen cell production and suppression of worker ovary activation. Here we show that the queen signal of the Brazilian stingless bee Friesella schrottkyi is a mixture of cuticular hydrocarbons. Stingless bees are therefore similar to ants, wasps and bumble bees, but differ from honey bees in which the queen's signal mostly comprises volatile compounds originating from the mandibular glands. This shows that cuticular hydrocarbons have independently evolved as the queen's signal across multiple taxa, and that the honey bees are exceptional. We also report the distribution of four active queen-signal compounds by Matrix-assisted laser desorption/ionization (MALDI) imaging. The results indicate a relationship between the behavior of workers towards the queen and the likely site of secretion of the queen's pheromones.

Nest thermoregulation in Polybia scutellaris (White) (Hymenoptera: Vespidae).

Polybia scutellaris (White) builds large nests characterized by numerous spiny projections on the surface. In order to determine whether or not the nest temperature is maintained because of homeothermic conditions of the nest individuals or otherwise, we investigated the thermal conditions within the nests built by P. scutellaris. We measured the temperature within active and abandoned nests. The temperature in the active nest was almost stable at 27°C during data collection, whereas the temperature in the abandoned nest varied with changes in ambient temperature. These results suggest that nest temperature was maintained by the thermogenesis of the individuals of the colony. This is the first report of nest incubation caused by thermogenesis of species of Polybia wasps.

The acceptance rate of young wasps by alien colonies depends on colony developmental stages in the swarm-founding wasp, Polybia paulista von ihering (Hymenoptera: Vespidae)

In social insects, newly emerged individuals learn the colony-specific chemical label from their natal comb shortly after their emergence. These labels help to identify each individual's colony of origin and are used as a recognition template against which individuals can discriminate nestmates from non-nestmates. Our previous studies with Polybia paulista von Ihering support this general pattern, and the acceptance rate of young female and male wasps decreased as a function of their age. Our study also showed in P. paulista that more than 90 percent of newly emerged female wasps might be accepted by conspecific unrelated colonies. However, it has not been investigated whether the acceptance rate of newly emerged female wasps depends on colony developmental stage of recipient colonies. We introduced newly emerged female wasps of P. paulista into different colony developmental stags of recipient colonies, i.e., worker-producing and male-producing colonies. We found that the acceptance rate of newly emerged female wasps by alien colonies was pretty lower by male-producing colonies than worker-producing colonies. This is the first study to show that the acceptance rate of young female wasps depends on stages of recipient colonies.

Nest thermoregulation in Polybia scutellaris (White) (Hymenoptera: Vespidae)

Polybia scutellaris (White) builds large nests characterized by numerous spiny projections on the surface. In order to determine whether or not the nest temperature is maintained because of homeothermic conditions of the nest individuals or otherwise, we investigated the thermal conditions within the nests built by P. scutellaris. We measured the temperature within active and abandoned nests. The temperature in the active nest was almost stable at 27°C during data collection, whereas the temperature in the abandoned nest varied with changes in ambient temperature. These results suggest that nest temperature was maintained by the thermogenesis of the individuals of the colony. This is the first report of nest incubation caused by thermogenesis of species of Polybia wasps.

Castes in the neotropical social wasp Leipomeles dorsata (Fabricius) (Hymenoptera: Vespidae): a window for workers achieving a new status in the colony.

Morphometric studies performed in several species of Neotropical social wasps from the tribe Epiponini showed that in some species there are marked differences between castes, while other species present highly distinct castes with differences only in ovarian development. This work analyzed females from six colonies of the social wasp Leipomeles dorsata (Fabricius) in which queens (egglayers) and workers showed differences in ovarian development and coloration. We propose that wasps with developed ovaries (egglayers) and coloration similar to those of workers are possibly intermediates that obtained the status of queens in the colony.

Castes in the neotropical social wasp Leipomeles dorsata (Fabricius) (Hymenoptera: Vespidae): a window for workers achieving a new status in the colony

Morphometric studies performed in several species of Neotropical social wasps from the tribe Epiponini showed that in some species there are marked differences between castes, while other species present highly distinct castes with differences only in ovarian development. This work analyzed females from six colonies of the social wasp Leipomeles dorsata (Fabricius) in which queens (egglayers) and workers showed differences in ovarian development and coloration. We propose that wasps with developed ovaries (egglayers) and coloration similar to those of workers are possibly intermediates that obtained the status of queens in the colony.

The acceptance rate of young wasps by alien colonies depends on colony developmental stages in the swarm-founding wasp, Polybia paulista von ihering (Hymenoptera: Vespidae).

In social insects, newly emerged individuals learn the colony-specific chemical label from their natal comb shortly after their emergence. These labels help to identify each individual's colony of origin and are used as a recognition template against which individuals can discriminate nestmates from non-nestmates. Our previous studies with Polybia paulista von Ihering support this general pattern, and the acceptance rate of young female and male wasps decreased as a function of their age. Our study also showed in P. paulista that more than 90% of newly emerged female wasps might be accepted by conspecific unrelated colonies. However, it has not been investigated whether the acceptance rate of newly emerged female wasps depends on colony developmental stage of recipient colonies. We introduced newly emerged female wasps of P. paulista into different colony developmental stags of recipient colonies, i.e., worker-producing and male-producing colonies. We found that the acceptance rate of newly emerged female wasps by alien colonies was pretty lower by male-producing colonies than worker-producing colonies. This is the first study to show that the acceptance rate of young female wasps depends on stages of recipient colonies.

Mandibular gland secretions of meliponine worker bees: further evidence for their role in interspecific and intraspecific defence and aggression and against their role in food source signalling.

Like ants and termites some species of stingless bees (Meliponini), which are very important pollinators in the tropics, use pheromone trails to communicate the location of a food source. We present data on the communicative role of mandibular gland secretions of Meliponini that resolve a recent controversy about their importance in the laying of such trails. Volatile constituents of the mandibular glands have been erroneously thought both to elicit aggressive/defensive behaviour and to signal food source location. We studied Trigona spinipes and Scaptotrigona aff. depilis ('postica'), two sympatric species to which this hypothesis was applied. Using extracts of carefully dissected glands instead of crude cephalic extracts we analysed the substances contained in the mandibular glands of worker bees. Major components of the extracts were 2-heptanol (both species), nonanal (T. spinipes), benzaldehyde and 2-tridecanone (S. aff. depilis). The effect of mandibular gland extracts and of individual components thereof on the behaviour of worker bees near their nest and at highly profitable food sources was consistent. Independent of the amount of mandibular gland extract applied, the bees overwhelmingly reacted with defensive behaviour and were never attracted to feeders scented with mandibular gland extract or any of the synthetic chemicals tested. Both bee species are capable of using mandibular gland secretions for intra- and interspecific communication of defence and aggression and share 2-heptanol as a major pheromone compound. While confirming the role of the mandibular glands in nest defence, our experiments provide strong evidence against their role in food source signalling.

Morphological studies on castes of Protopolybia chartergoides (Hymenoptera, Vespidae, Epiponini) observed in colonies during male production stage

Social wasps are important for the study of social behavior evolution because their colonies present different degrees of caste differentiation, from morphologically similar to highly conspicuous, associated with the occasional presence of intermediate females, which bears developed ovaries but no insemination. In the Polistinae, depending on the taxon, such differentiation can be discreet or conspicuous. This work intended to study morphological and physiological differences between castes in Protopolybia chartergoides by using morphometric analyses associated with multivariate statistical analyses and physiological evaluations from females' ovarian development. Results evidence low morphological and physiological differences among the castes in P. chartergoides, indicating three groups of females: queens, workers and intermediates. In this way, it was possible to suggest that Protopolybia chartergoides presents post-imaginal caste differentiation (or a very subtle form of pre-imaginal determination).

Vespas sociais são importantes para o estudo da evolução do comportamento social devido suas colônias apresentarem diferentes graus de diferenciação de castas, de morfologicamente similares às altamente conspícuas, associadas com a presença de fêmeas intermediárias, as quais desenvolvem seus ovários, mas não estando inseminadas. Nos Polistinae, tais diferenciações podem ser discretas ou conspícuas. Este trabalho teve como objetivo estudar as diferenças morfológicas e fisiológicas entre as castas de Protopolybia chartergoides pelo uso de análises morfométricas associada com análises estatísticas multivariadas e avaliações fisiológicas do desenvolvimento ovariano das fêmeas. Os resultados mostraram baixa diferenciação morfológica e fisiológica entre as castas de P. chartergoides, indicando a presença de três grupos de fêmeas: rainhas, operárias e intermediárias. Assim, foi possível sugerir que Protopolybia chartergoides apresenta diferenciação de casta pós-imaginal (ou uma forma muito sutil de determinação pré-imaginal).

Castes and asynchronous colony cycle in Polybia bistriata (Fabricius) (Hymenoptera: Vespidae)

Neotropical swarm-founding wasps, the Epiponini, are an outstanding group of social insects whose societies are polygynic and complex nest builders. Caste dimorphism in these wasps ranges from incipient to highly distinct. Morphometric analyses of nine body parts, ovarian status, relative age and development of the 5th gastral sternite gland (Richards´ gland) of Polybia bistriata Fabricius were undertaken in order to estimate caste differentiation in nine colonies. Morphological and physiological data were used in multivariate analyses in order to evaluate the level of discrimination between inseminated and non-inseminated females. Clear physiological differences were found: queens had highly developed ovaries and they were inseminated, and workers had totally undeveloped ovaries or they had few developed oocytes (only in two colonies), but in both cases insemination was not detected. ANOVA and discriminant function analysis detected slight, but significant differences between castes. In relation to colony cycle, colonies were considered to be in the following stages: one in pre-emergence, four in worker production and four in sexuals production. Richards´ gland analyses indicated that in small colonies (<100 females) queens had a less developed gland than in medium (100-200 females), and large colonies (>200 females). Taking the whole data, it was possible to conclude that caste differences were slight, but more evident in some phases of the colony cycle, a phenomenon previously described for other epiponines.

Os Epiponini, vespas sociais neotropicais, são um grupo de insetos sociais com sociedades poligínicas, construtoras de ninhos complexos. O dimorfismo das castas varia de incipiente até altamente distinto. Análises morfométricas de nove partes corporais, estágio de desenvolvimento ovariano, idade relativa e desenvolvimento da glândula do quinto esternito do gáster (glândula de Richards) de Polybia bistriata (Fabricius) foram utilizados para calcular o nível de diferenciação das castas de nove colônias. Dados morfológicos e fisiológicos foram usados em análises multivariadas para avaliar o nível de discriminação entre fêmeas inseminadas e não-inseminadas. Foram encontradas diferenças fisiológicas claras: rainhas tinham ovários altamente desenvolvidos e inseminados, e as operárias não estavam inseminadas e tinham ovários não desenvolvidos ou uns poucos oócitos desenvolvidos (em duas colônias). ANOVA e análise da função discriminante detectaram diferenças significantes entre as castas. Considerou-se, pelos dados obtidos, que as colônias estavam nas seguintes fases em relação ao ciclo colonial: uma em pré-emergência, quatro em produção de operárias ;.e quatro em produção de sexuados. Análises das glândulas de Richards indicaram o seu menor desenvolvimento em rainhas de colônias pequenas (< 100 fêmeas) do que aquelas de colônias médias (100-200 fêmeas), e grandes (> 200 fêmeas). Considerando-se todos os dados, é possível concluir que as diferenças entre as castas são pequenas, mas evidentes em algumas fases do ciclo colonial, um fenômeno previamente descrito para outros epiponíneos.

Castes and asynchronous colony cycle in Polybia bistriata (Fabricius) (Hymenoptera: Vespidae).

Neotropical swarm-founding wasps, the Epiponini, are an outstanding group of social insects whose societies are polygynic and complex nest builders. Caste dimorphism in these wasps ranges from incipient to highly distinct. Morphometric analyses of nine body parts, ovarian status, relative age and development of the 5th gastral sternite gland (Richards' gland) of Polybia bistriata Fabricius were undertaken in order to estimate caste differentiation in nine colonies. Morphological and physiological data were used in multivariate analyses in order to evaluate the level of discrimination between inseminated and non-inseminated females. Clear physiological differences were found: queens had highly developed ovaries and they were inseminated, and workers had totally undeveloped ovaries or they had few developed oocytes (only in two colonies), but in both cases insemination was not detected. ANOVA and discriminant function analysis detected slight, but significant differences between castes. In relation to colony cycle, colonies were considered to be in the following stages: one in pre-emergence, four in worker production and four in sexuals production. Richards' gland analyses indicated that in small colonies (<100 females) queens had a less developed gland than in medium (100-200 females), and large colonies (>200 females). Taking the whole data, it was possible to conclude that caste differences were slight, but more evident in some phases of the colony cycle, a phenomenon previously described for other epiponines.

Non-ovipositing nurse workers with developed ovaries in trigona cilipes cilipes (Hymenoptera, Meliponini)

Numerous species of tropical stingless bees (Hymenoptera, Meliponini) have workers that lay eggs even inthe presence of a queen, i.e., queenright colonies. These worker eggs, which are laid during cell provisioningand oviposition process (POP), are eaten mainly by the queen. In this work, we studied POP in a colonyof Trigona cilipes in the queenright and queenless stages. POP behavior was observed on 67 occasions. Inaddition, the ovaries of different aged workers in both stages were analyzed. Although workers in both stagesof the colony possessed developed ovaries, they never oviposited, despite showing behavior similar to thatseen in species with workers which lay eggs. This form of sterility is outstanding among social insects andis comparable only to other meliponines that have fully sterile workers.

Males of Nectarinella Bequaert (Hymenoptera, Vespidae, Polistinae)

Os machos das duas espécies de vespas sociais do gênero Nectarinella Bequaert, N. xavantinensis Mateus & Noll e N. championi (Dover), são descritos e suas genitálias ilustradas. São feitas comparações com espécies de gêneros estreitamente relacionados, Chartergellus, Leipomeles, Marimbonda, Parachartergus e Pseudopolybia.

Predatory behavior in a necrophagous bee Trigona hypogea (Hymenoptera; Apidae, Meliponini).

Although most bees feed on nectar and pollen, several exceptions have been reported. The strangest of all is the habit found in some neotropical stingless bees, which have completely replaced pollen-eating by eating animal protein from corpses. For more than 20 years, it was believed that carrion was the only protein source for these bees. We report that these bees feed not only off dead animals, but on the living brood of social wasps and possibly other similar sources. Using well developed prey location and foraging behaviors, necrophagous bees discover recently abandoned wasps' nests and, within a few hours, prey upon all immatures found there.