Nest Architecture Development of Grass-Cutting Ants, Atta capiguara (Hymenoptera: Formicidae)

ABSTRACT Atta capiguara grass-cutting ants are commonly found in the Cerrado biome, in open fields. Although grass-cutting ants build giant nests, little has been elucidated about this building pattern and when chambers and tunnels emerge. The present study describes the nest architecture development of A. capiguara grass-cutting ants from data on 31 cement-molded nests. A. capiguara nests grow with increases in the number of fungus chambers and emergence and increase of waste chambers and foraging tunnels. The structural growth of A. capiguara nests in the first year and a half of age (18 months) is vertical, with the building of the first chambers in the soil profile. After 18 months, the nests grow sideways with the addition of chambers and tunnels, and the first waste chambers appear. Between 18 and 54 months, the number of fungus chambers increases from 1-3 to 21-32, and the chambers are concentrated at the soil surface, although they can be found more than 3 m deep. In addition, the total volume of the waste chambers increases with the increment in the fungus chambers volume. Thus, this study contributes to understanding the nest architecture development of A. capiguara grass-cutting ants and demonstrates that the total volume of waste chambers is proportional to the total volume of fungus chambers suitable for the colony.

Effect of the presence of brood and fungus on the nest architecture and digging activity of Acromyrmex subterraneus Forel (Hymenoptera, Formicidae)

ABSTRACT This study investigated the stimuli that trigger digging behavior in Acromyrmex subterraneus during nest building. The hypothesis was that the presence of the fungus garden and/or brood triggers the excavation of tunnels and chambers. For the experiment, the excavation rate of individually marked workers kept in plastic cylinders filled with soil was recorded. Four treatments were applied: (1) 30 medium-sized workers, 5 g fungus garden and 30 brood items (larvae and pupae); (2) 30 medium-sized workers and 5 g fungus garden; (3) 30 medium-sized workers and 30 brood items; (4) 30 medium-sized workers without fungus and brood. After 24 h, morphological parameters of nest structure (length and width of the chambers and tunnels in cm) and the volume of excavated soil were recorded. In contrast to the expected findings, no change in morphological structure, rate of excavation by workers, or volume of excavated soil was observed between treatments, except for tunnel width, which was greater, when no brood or fungus garden was present. Thus, the results do not support the hypothesis that the fungus garden and/or brood are local stimuli for nest excavation or that they mold the internal architecture of the nest. Although this hypothesis was confirmed for Acromyrmex lundii and Atta sexdens rubropilosa, the same does not apply to A. subterraneus. The digging behavior of workers is probably the result of adaptation during nest building in different habitats.

Efeito de grupo versus produção de CO2 em operárias da formiga cortadeira Atta sexdens rubropilosa / Effect of group versus CO2 production in the leaf cutting ant workers Atta sexdens rubropilosa

Será que a vida em grupo acarreta uma maior atividade individual, e consequentemente, um maior gasto energético? Para responder a essa questão, hipotetizou-se que exista uma maior produção de CO2, quando se aumenta o tamanho do grupo de formigas operárias e, consequentemente, um maior custo energético ao indivíduo quando está em grupo. Assim, grupos de 10, 20, 30, 40 e 50 operárias foram fechados em uma câmara hermética, durante 24 horas. Posteriormente, efetuou-se a medição das concentrações de CO2 nos recipientes respirométricos. Ao contrário do esperado, a produção de CO2, e consequentemente, o gasto energético individual, não diferiram estatisticamente quando se aumentou o tamanho do grupo de operárias. Dessa forma, refuta-se a hipótese de que o tamanho do grupo conduz a um maior custo energético individual, haja vista a maior interação entre os indivíduos. O estudo com operárias de Atta sexdens rubropilosa evidenciou que o tamanho do grupo não conduz a um maior custo energético individual, sendo a produção de CO2 e, consequentemente, o gasto energético individual similar, independente do tamanho do grupo.

Does the social life entail greater individual activity, and consequently, higher energy expenditure? To answer this question, we hypothesized that there is higher CO2 production, when we increase the size of the group of workers, and hence a higher energy cost to the individual when they are in groups. Thus, groups of 10, 20, 30, 40 and 50 workers were sealed in a hermetic chamber for 24 hours. Subsequently, we performed the measurements of the CO2 concentration in the containers respirometric. Unlike the expected CO2 production, and consequently the individual energy expenditure did not differ when we increase the size of the group of workers. Thus, we refuted the hypothesis that the group size leads to a higher cost individual energy, since the greater interaction between individuals. In conclusion, our study with Atta sexdens rubropilosa workers determined that the size of the group does not lead to higher energy costs individual and CO2 production, and therefore energy expenditure similar individual, independent of the group size.

An adaptive view of caste differentiation in the neotropical wasp Polybia (Trichothorax) sericea Olivier (Hymenoptera: Vespidae)

The tribe Epiponini comprehends the swarm-founding Neotropical wasps, with several species endemic to Brazil, which are extremely important in studies of social evolution of wasps. The Epiponini diverge in several ways from the definitions of high eusociality, since caste syndromes range from species without morphological caste differentiation to those with complete caste dimorphism, and all species are polygynous. Frequently, indirect studies based on morphometry and physiology are the only solutions to collect data regarding the natural history and caste system in this tribe, since most species are extremely aggressive and build enveloped nests, usually in places of difficult access. We analyzed morphological parameters in seven colonies of the Epiponini species Polybia (Trichothorax) sericea Olivier in different phases of colonial development. Nine body variables were taken and females were classified according to their ovary development and spermathecal contents. The results showed that caste differences in this species are based on a contrast among variables: queens have larger mesosoma and abdomen, but are smaller in head width and wing length. These results suggest that morphological caste differentiation in this species is based mainly on body shape. We considered this combination of characters as being adaptive. We also showed that caste differences varied according to the colony cycle, with more conspicuous differences when queen number is reduced.

Sequence and expression pattern of the germ line marker vasa in honey bees and stingless bees

Queens and workers of social insects differ in the rates of egg laying. Using genomic information we determined the sequence of vasa, a highly conserved gene specific to the germ line of metazoans, for the honey bee and four stingless bees. The vasa sequence of social bees differed from that of other insects in two motifs. By RT-PCR we confirmed the germ line specificity of Amvasa expression in honey bees. In situ hybridization on ovarioles showed that Amvasa is expressed throughout the germarium, except for the transition zone beneath the terminal filament. A diffuse vasa signal was also seen in terminal filaments suggesting the presence of germ line cells. Oocytes showed elevated levels of Amvasa transcripts in the lower germarium and after follicles became segregated. In previtellogenic follicles, Amvasa transcription was detected in the trophocytes, which appear to supply its mRNA to the growing oocyte. A similar picture was obtained for ovarioles of the stingless bee Melipona quadrifasciata, except that Amvasa expression was higher in the oocytes of previtellogenic follicles. The social bees differ in this respect from Drosophila, the model system for insect oogenesis, suggesting that changes in the sequence and expression pattern of vasa may have occurred during social evolution.

Morphological caste studies in the neotropical swarm-founding Polistinae wasp Angiopolybia pallens (Lepeletier) (Hymenoptera: Vespidae) / Estudos morfológicos das castas da vespa Polistinae enxameadora neotropical, Angiopolybia pallens (Lepeletier) (Hymenoptera: Vespidae)

Neotropical swarm-founding wasps present polygynic colonies and a great variation in relation to caste differentiation, ranging from species in which queens and workers are similar in shape and size to those where variation in shape and size is conspicuous. Canonical discriminant analysis and Bonferroni t-test analysis on morphometric data collected from eight body parts of specimens from nine colonies of Angiopolybia pallens (Lepeletier) were undertaken as a step towards to a better understanding on caste differenciation in Epiponini. All specimens were dissected to verify the ovary developmental stage, and the spermatheca was removed to check for the mating status. Females were then grouped as queens (mated with developed ovaries), intermediates (virgin with developed ovaries) and workers (virgin with undeveloped ovaries). Even though differences were found for some measurements in seven out of the nine colonies evaluated, multivariate analysis (Wilks' lambda values) indicated castes could not be reliably distinguished only by morphology. We conclude that morphological differences among A. pallens castes are low or absent, reflecting a possible groundplan for the Epiponini, i.e. a post-imaginal pattern of caste differentiation.

As vespas neotropicais enxameadoras possuem colônias poligínicas e uma variação muito grande em relação à diferenciação das castas, desde espécies em que rainhas e operárias possuem formas e tamanhos semelhantes até aquelas em que grande diferença na forma e no tamanho são observados. Estudos morfométricos utilizando oito medidas corporais em nove colônias de Angiopolybia pallens (Lepeletier) foram realizados a fim de aumentar o conhecimento sobre diferenciação de castas em Epiponini, empregando-se análise canônica discriminante e teste t de Bonferroni. Os indivíduos foram dissecados para determinar o nível de desenvolvimento ovariano e as espermatecas foram removidas para verificar a ocorrência de inseminação e assim definir os três grupos de fêmeas: rainhas (ovários desenvolvidos e espermateca fecundada), intermediárias (ovários desenvolvidos e espermateca não-fecundada) e operárias (ovários não-desenvolvidos). Apesar de serem encontradas diferenças significativas para algumas medidas em sete das nove colônias, a análise multivariada (lambda de Wilks) mostrou que as castas são de difícil distinção apenas pela morfologia. É possível concluir que diferenças morfológicas entre castas de A. pallens são pequenas ou ausentes, o que reflete o plano basal para os Epiponini, i.e. padrão pós-imaginal de diferenciação castas.

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.