As vespas da tribo Epiponini apresentam colônias com várias rainhas, cujo número varia durante o ciclo colonial. Nas colônias há inicialmente um número maior de rainhas, mas conforme o ciclo se desenvolve seu número diminui. Tem sido argumentado que nas colônias com indivíduos totipotentes pode haver conflitos de interesse entre rainhas e operárias. Neste contexto as operárias que são maioria poderiam ter uma vantagem para regular a produção de rainhas. Nos Epiponini o controle reprodutivo é feito possivelmente por sinalização química, que junto com posturas ritualizadas comunicam a presença da rainha na colônia. Como o processo de produção de rainhas é pouco conhecido, o objetivo deste trabalho foi estudar o comportamento de rainhas e operárias durante a fase de produção e seleção de modo a interpretar filogeneticamente as mudanças comportamentais entre as espécies. O estudo foi conduzido utilizando-se nove espécies de cinco gêneros: Chartergellus, Metapolybia, Protopolybia, Polybia, Brachygastra. Em cada colônia foram marcadas todas as rainhas, algumas operárias, e fêmeas recém-emergidas para realizar observações focais e filmagens dos comportamentos. Para promover a produção de rainhas foram feitos testes de remoção. Alguns dos comportamentos característicos das rainhas, e aqueles associados ao processo de seleção, tiveram sua origem interpretada filogeneticamente. O comportamento que melhor identifica as rainhas é o bending display I (BD1), um display de dominância feito das rainhas para as operárias, que foi basal para os Epiponini. O BD1 feito das operárias para as rainhas é uma apomorfía das operárias de Metapolybia. Outros comportamentos de teste entre rainhas surgiram independentemente em algumas espécies e poderiam estar relacionados com um aumento na complexidade das interações. Os testes das operárias estariam presentes no ancestral dos Epopinini indicando que elas controlam a seleção das rainhas. O comportamento de worker dance é utilizado principalmente pelas operarias dos clados de Polybia e Metapolybia para testar a capacidade reprodutiva das rainhas. Parece que os comportamentos ritualizados de dominância e de teste são utilizados como sinais honestos do potencial reprodutivo das fêmeas em vez de comportamentos agressivos. Em Metapolybia e Protopolybia exigua o BD1 é realizado com maior frequência após a eliminação da rainha e durante a seleção, diferente de Chartergellus e Protopolybia picteti onde não houve mudança. Em Chartergellus não houve um aumento dos displays, porque as fêmeas no geral não interagiam e a seleção é feita mediante a expulsão das rainhas. Em P. picteti nem todas as rainhas foram eliminadas. Em Metapolybia e Chartergellus no início do ciclo colonial há tolerância reprodutiva, e a eliminação das rainhas estimulou muitas fêmeas a ovipositar, mas o número eventualmente diminui. Este resultado mostra que a delimitação de casta poderia ser feita mediante algum tipo de sinalização química. Em Protopolybia há pouca redução do número de rainhas ao longo do ciclo e as intermediárias constantemente tentam botar ovos. Para as intermediárias é conveniente manter os ovários ativos para tentar atingir o status de rainha em uma nova colônia. A totipotência das fêmeas para desenvolver seus ovários é basal para os Epiponini. As operárias e intermediárias jovens, de idade intermediária e velhas podem desenvolver seus ovários diante da ausência da rainha. A flexibilidade é um traço muito importante nos Epiponini, uma vez que permite à população responder a situações adversas, mobilizando indivíduos a realizar diferentes tarefas segundo as necessidades da população. / Epiponini wasps have colonies with multiple queens, which alternate their number under different stages of the colony cycle. Initially there are several potential queens, but as the colony cycle develops, queen number is reduced. Because most of individuals remain reproductively totipotent there is great potential for conflicts over reproduction. Under this conditions workers could have an advantage to control queen production because they are the dominant caste. Reproductive control could be performed by chemical signals, which along with ritualized behaviors communicate queen presence in the colony. However, queen selection process is a little known across Epiponini, and for this reason our goal was to study queens and workers behavior during queen selection phase, performing a phylogenetic analysis to understand behavioral changes among species. We performed observations in nine species belong to five genera: Chartergellus, Metapolybia, Protopolybia, Polybia, Brachygastra. We marked queens, a sample of workers and newly emerged females to perform direct and video focal observations. To induce queen production, we performed queen removal tests. Some of queens characteristic behaviors and those associated with queen selection process were analyzed via phylogenetic optimization. Bending display I (BD1) is a dominance display performed queens against workers, is the behavior that better identify queens and is also basal for all Epiponini clades. BD1 perform from workers to queens is an apomorphy of Metapolybia. Another test behaviors performed among queens appeared independently in some species, and could be related with an increase of interaction complexity. Worker tests were present in the Epiponini ancestor, indicating that workers are in charge of queen control. Worker dance behavior was synapomorphic for the clade composed by Polybia, Metapolybia and their close related genera, in which workers test queens reproductive potential. Apparently, ritualized tests and dominance behaviors are used in Epiponini as honest signals of queens reproductive potential instead of aggressive behaviors. In Metapolybia and in Protopolybia exigua, BD1 is performed with high frequency after queen elimination and during queen selection process. On the other hand, in Chartergellus and Protopolybia picteti displays frequency did not change. In Chartergellus BD1 frequency did not change because females do not interact to each other a lot, also queen selection is performed by queens expulsion instead by tests. In P. picteti we possibly failed to remove all queens from the colony. Metapolybia and Chartergellus have reproductive tolerance between females during first stages of colony cycle, and queen elimination induced females to lay eggs, but queen number eventually is reduced. These results suggest that reproductive control in fact could be performed by chemical signals. On the other hand, in Protopolybia queens number change little during the colony cycle. In adition, intermediates constantly attempt to lay eggs. Persistence oviposition could be convenient for intermediates in Protopolybia to achieve queen status in new colonies. Females totipotency to develop ovaries is also basal for Epiponini. Not only young females develop their ovaries under queen lost; old and middle age individuals also attempt to replace the queen. Caste flexibility is decisive for colony survival of swarm wasps because allows colonies to respond efficiently to different situations that may eventually arise.

A região noroeste do Estado de São Paulo é uma das que mais sofre com o processo de fragmentação da vegetação natural que foi substituída principalmente por ambientes urbanizados, pastagens e áreas agriculturáveis. Cana-de-açúcar e citricultura são duas das principais monoculturas presentes atualmente na região. A área de entorno (matriz) dos fragmentos florestais exerce influência nas populações que habitam seu interior, incluindo o grupo das vespas sociais. O presente trabalho teve o objetivo de realizar um levantamento das vespas sociais (Polistinae) presentes em fragmentos florestais do Noroeste Paulista com matrizes de entorno compostas por canaviais e laranjais. Utilizamos uma metodologia baseada em coleta ativa utilizando líquido atrativo em um transecto no interior e um na borda dos fragmentos e obtivemos 20 espécies em Magda, 13 em Bebedouro, 13 em Matão e 19 em Barretos. Verificamos uma maior riqueza de espécies na borda dos fragmentos. O gênero mais representativo foi Agelaia em todas as áreas analisadas. Os maiores índices de diversidade de Shannon-Wiener foram obtidos nos transectos da borda de Barretos (H=2,26) e borda de Magda (H=2,25), fragmentos circundados por laranja e cana-de-açúcar respectivamente. Analisando a área como umtodo, Magda apresentou a maior diversidade (H=2,12). Espécies como Brachygastra moebiana, Metapolybia docilis, Mischocyttarus ignotus, M. paulistanus e M. consimilis não haviam sido registrados em levantamentos recentes no estado, além disso o registro de M. consimilis aparentemente é novo para o estado de São Paulo. Realizamos comparações com outros levantamentos realizados no estado e verificamos um indício de que a riqueza de vespas sociais pode estar relacionada com a quantidade de vegetação natural remanescente da área. Concluímos que com nossos dados não foi possível observar uma relação entre a ocorrência de vespas sociais e a matriz de entorno. / The northwest of São Paulo State is a region that undergoes a fragmentation process of its natural vegetation that was replaced by urban areas, meadows or farming lands. Sugarcane and citrus are two of the main regional monocultures. The surrounding area (matrix) of forest fragments influences the populations that inhabit the interior, including social wasps. The present work aimed at surveying social wasps (Polistinae) present in forest fragments of northwest of São Paulo State with different surrounding matrices composed by sugarcane and citrus. We used a methodology based on active collection using and attractive liquid in one transect on the edge and one on the interior of the fragments and we collected 20 species in Magda, 13 in Bebedouro, 13 in Matão and 19 in Barretos. We noted a greater richness on the edge of the fragments. Agelaia was the most representative genus in all of the areas. The greatest indices of Shannon-Wiener diversity were obtained on the edge of Barretos (H=2,26) and on the edge of Magda (H=2,25), fragments surrounded by citrus and sugarcane respectively. If the total area was analyzed, Magda showed the greater diversity (H=2,12). Species such as Brachygastra moebiana, Metapolybia docilis, Mischocyttarus ignotus, M. paulistanus and M. consimilis had not been recorded on recent surveys on the state, furthermore Mischocyttarus consimilis apparently is a new record for the São Paulo State. We did comparisons with others surveys made on the state and we noted that social wasps richness can be related with the amount of remnants of natural vegetation in a given area. We concluded that with our data it was not possible to observe a relation between the occurrence of social wasps and the surrounding matrix.

Submitted by AGDA ALVES DA ROCHA null (rocha.agda@gmail.com) on 2017-11-28T19:58:09Z No. of bitstreams: 1 Rocha_Agda_2017.pdf: 3754795 bytes, checksum: 147e7d00c8330084beb9e117e77ac336 (MD5) / Approved for entry into archive by LUCIA SILVA PARRA null (lucia@ia.unesp.br) on 2017-11-29T16:34:46Z (GMT) No. of bitstreams: 1 rocha_aa_dr_rcla.pdf: 3754795 bytes, checksum: 147e7d00c8330084beb9e117e77ac336 (MD5) / Made available in DSpace on 2017-11-29T16:34:46Z (GMT). No. of bitstreams: 1 rocha_aa_dr_rcla.pdf: 3754795 bytes, checksum: 147e7d00c8330084beb9e117e77ac336 (MD5) Previous issue date: 2017-03-13 / A vespa social basal Mischocyttarus nomurae Richards tem ocorrência exclusiva no Brasil e foi registrada nos estados do Ceará, Bahia e Minas Gerais. Na Bahia, ocorre em três municípios da Chapada Diamantina: Lençóis, Mucugê e Rio de Contas. Nesta última, foram estudadas populações com relação à sua biologia básica, buscando responder perguntas relacionadas aos seguintes aspectos: morfologia externa dos imaturos, perfis comportamental, morfofisiológico e químico, substituição de rainhas, arquitetura de ninhos e hábitos de nidificação. Com relação à morfologia dos imaturos, a taxa média de crescimento da larva foi de 1,48. Com exceção da larva de 1o ínstar, que revelou-se menor do que o ovo, a espécie apresentou o padrão do grupo: presença de lobos, estes em número de dois e projetados para a frente, só totalmente desenvolvidos quando a larva atinge o 5o instar; as mandíbulas das larvas de 5o instar apresentam um único dente alongado e o primeiro espiráculo é mais do que duas (3,1) vezes maior em diâmetro que os demais. As cápsulas cefálicas da larva de 5º ínstar e da pré-pupa não diferiram significativamente. Após os estudos dos perfis morfofisiológico, químico e comportamental, foi verificado que as fêmeas de M. nomurae não apresentam distinção em sua morfologia externa, mas cinco tipos de desenvolvimento ovariano foram observados, três menos desenvolvidos (A1, A2 e B) e dois mais desenvolvidos (C e D). Foram detectados 34 compostos na sua cutícula, cujas cadeias variaram de 18 a 33 átomos de carbono. Os compostos consistiram em alcanos lineares (9%), alcanos ramificados (89%) e alcenos (2%). Foram identificados três grupos de fêmeas: Operária 1, Operária 2 e Rainha e a análise discriminante do perfil dos hidrocarbonetos cuticulares relacionada a esses grupos de fêmeas apresentou um p-valor significativo (Wilks' lambda= 0,135, F= 1.665, p = 0,0227). Após oito dias da remoção da fêmea dominante do ninho, em todas as colônias, exceto uma, a fêmea era anteriormente a fêmea β. Fêmeas com ovários não desenvolvidos podem realizar comportamentos de fêmea dominante, assim como uma fêmea com ovário desenvolvido comportouse como forrageadora típica. Houve diferença entre algumas atividades exercidas pelas fêmeas posicionadas em 2º lugar no ranking (fêmea β) e as demais fêmeas do ninho (3ª posição em diante), antes e após a remoção experimental da rainha. M. nomurae apresenta um sistema de gerontocracia flexível na sua substituição de rainhas e suas fêmeas sucessoras tornam-se mais agressivas que as rainhas originais, assumindo tarefas de dominantes, mesmo sem estarem inseminadas. Como algumas espécies do gênero, a maioria das colônias apresentou hábitos de nidificação relacionados a ambientes antrópicos. Destes, 61,2% das colônias utilizaram material de origem vegetal (madeira e palha). A altura média de construção do ninho com relação ao solo foi elevada, semelhante as das demais espécies do gênero (2,20 ± 0,83, 0,45 – 5,00m). A maioria dos ninhos foi construída com o favo voltado para baixo (70,15%) e em substrato horizontal (53,73%). O único favo do ninho pode ser arredondado ou elíptico, raramente retangular. O número de camadas de mecônio variou de zero a cinco e a espécie M. nomurae apresenta ninhos pequenos, com suas colônias apresentando um baixo número de adultos produzidos, embora haja uma alta taxa de reutilização de células. / The basal social wasp Mischocyttarus nomurae Richards occurs exclusively in Brazil, having been recorded there in the states of Ceará, Bahia, and Minas Gerais. It is known to occur in three municipalities in the Chapada Diamantina mountains of Bahia State: Lençóis, Mucugê, and Rio de Contas. Population studies were undertaken in the latter municipality to address questions related to the basic biology of M. nomurae: the external morphologies of the immatures; their behavior; their morpho-physiological and chemical profiles; queen substitutions; nest architecture, and nidification habits. In terms of the morphologies of the immatures, the mean larval growth rate was 1.48. With the exception of the 1st instar larvae, which are smaller than the eggs, the species demonstrated the standard developmental pattern for the group: the presence of lobes (two) projected forward, those only becoming fully developed in the 5th instar stage; mandibles in the 5th instar stage with a single elongated tooth, and the first spiracle with a diameter more than two (3.1) times greater than the others. The cephalic capsules of the 5th instar larvae and pre-pupae do not significantly differ. Studies of the morpho-physiological, chemical, and behavioral profiles of those wasps indicated that the females of M. nomurae do not have distinct external morphologies, although five different types of ovarian development were observed: three less-developed types (A1, A2 and B); and two more developed types (C and D). Thirty-four distinct chemical compounds were detected in the cuticle, with carbon chains varying from 18 to 33 atoms. Those carbon compounds consisted of linear alkanes (9%), branched alkanes (89%), and alkenes (2%). Three groups of females were identified: Workers 1, Workers 2, and Queens. Discriminant analyses of the cuticular hydrocarbons of those female groups demonstrated a significant p-value (Wilks' lambda= 0.135, F= 1.665, p = 0.0227). Eight days after the removal of the dominant female from the nest, female substitutions were observed all of the colonies (except one) assuming the condition of a new queen (from the ranks of the β females). Females with undeveloped ovaries do not perform behaviors associated with dominant females, nor do females with developed ovaries behave as typical foragers. There were differences between some of the activities performed by second-rank females (β females) and other females in the same nest (from the 3rd position down), both before and after experimentally removing the queen. M. nomurae demonstrates a flexible gerontocratic system in terms of queen substitutions, and the new queens become more aggressive than the original queens, assuming dominant roles even without being inseminated. As with many species of the same genus, most of the colonies nidify in anthropically modified environments. Of those colonies, 61.2% built nests using plant material (wood and leaves). The nests were constructed at reasonably high mean heights above ground level (2.20 ± 0.83 m; from 0.45 – 5.00 m), similar to other species of the genus. Most of the nests were affixed to horizontal substrates (53.73%) and constructed with the comb facing downward (70.15%). A nest comb can be rounded or elliptic, rarely rectangular. The numbers of meconium layers varied from zero to five. The nests of M. nomurae are generally small, with their colonies producing only small numbers of adults - although with high rates of comb cell reutilization.

A região noroeste do Estado de São Paulo é uma das que mais sofre com o processo de fragmentação da vegetação natural que foi substituída principalmente por ambientes urbanizados, pastagens e áreas agriculturáveis. Cana-de-açúcar e citricultura são duas das principais monoculturas presentes atualmente na região. A área de entorno (matriz) dos fragmentos florestais exerce influência nas populações que habitam seu interior, incluindo o grupo das vespas sociais. O presente trabalho teve o objetivo de realizar um levantamento das vespas sociais (Polistinae) presentes em fragmentos florestais do Noroeste Paulista com matrizes de entorno compostas por canaviais e laranjais. Utilizamos uma metodologia baseada em coleta ativa utilizando líquido atrativo em um transecto no interior e um na borda dos fragmentos e obtivemos 20 espécies em Magda, 13 em Bebedouro, 13 em Matão e 19 em Barretos. Verificamos uma maior riqueza de espécies na borda dos fragmentos. O gênero mais representativo foi Agelaia em todas as áreas analisadas. Os maiores índices de diversidade de Shannon-Wiener foram obtidos nos transectos da borda de Barretos (H=2,26) e borda de Magda (H=2,25), fragmentos circundados por laranja e cana-de-açúcar respectivamente. Analisando a área como umtodo, Magda apresentou a maior diversidade (H=2,12). Espécies como Brachygastra moebiana, Metapolybia docilis, Mischocyttarus ignotus, M. paulistanus e M. consimilis não haviam sido registrados em levantamentos recentes no estado, além disso o registro de M. consimilis aparentemente é novo para o estado de São Paulo. Realizamos comparações com outros levantamentos realizados no estado e verificamos um indício de que a riqueza de vespas sociais pode estar relacionada com a quantidade de vegetação natural remanescente da área. Concluímos que com nossos dados não foi possível observar uma relação entre a ocorrência de vespas sociais e a matriz de entorno. / The northwest of São Paulo State is a region that undergoes a fragmentation process of its natural vegetation that was replaced by urban areas, meadows or farming lands. Sugarcane and citrus are two of the main regional monocultures. The surrounding area (matrix) of forest fragments influences the populations that inhabit the interior, including social wasps. The present work aimed at surveying social wasps (Polistinae) present in forest fragments of northwest of São Paulo State with different surrounding matrices composed by sugarcane and citrus. We used a methodology based on active collection using and attractive liquid in one transect on the edge and one on the interior of the fragments and we collected 20 species in Magda, 13 in Bebedouro, 13 in Matão and 19 in Barretos. We noted a greater richness on the edge of the fragments. Agelaia was the most representative genus in all of the areas. The greatest indices of Shannon-Wiener diversity were obtained on the edge of Barretos (H=2,26) and on the edge of Magda (H=2,25), fragments surrounded by citrus and sugarcane respectively. If the total area was analyzed, Magda showed the greater diversity (H=2,12). Species such as Brachygastra moebiana, Metapolybia docilis, Mischocyttarus ignotus, M. paulistanus and M. consimilis had not been recorded on recent surveys on the state, furthermore Mischocyttarus consimilis apparently is a new record for the São Paulo State. We did comparisons with others surveys made on the state and we noted that social wasps richness can be related with the amount of remnants of natural vegetation in a given area. We concluded that with our data it was not possible to observe a relation between the occurrence of social wasps and the surrounding matrix.

As vespas da tribo Epiponini apresentam colônias com várias rainhas, cujo número varia durante o ciclo colonial. Nas colônias há inicialmente um número maior de rainhas, mas conforme o ciclo se desenvolve seu número diminui. Tem sido argumentado que nas colônias com indivíduos totipotentes pode haver conflitos de interesse entre rainhas e operárias. Neste contexto as operárias que são maioria poderiam ter uma vantagem para regular a produção de rainhas. Nos Epiponini o controle reprodutivo é feito possivelmente por sinalização química, que junto com posturas ritualizadas comunicam a presença da rainha na colônia. Como o processo de produção de rainhas é pouco conhecido, o objetivo deste trabalho foi estudar o comportamento de rainhas e operárias durante a fase de produção e seleção de modo a interpretar filogeneticamente as mudanças comportamentais entre as espécies. O estudo foi conduzido utilizando-se nove espécies de cinco gêneros: Chartergellus, Metapolybia, Protopolybia, Polybia, Brachygastra. Em cada colônia foram marcadas todas as rainhas, algumas operárias, e fêmeas recém-emergidas para realizar observações focais e filmagens dos comportamentos. Para promover a produção de rainhas foram feitos testes de remoção. Alguns dos comportamentos característicos das rainhas, e aqueles associados ao processo de seleção, tiveram sua origem interpretada filogeneticamente. O comportamento que melhor identifica as rainhas é o bending display I (BD1), um display de dominância feito das rainhas para as operárias, que foi basal para os Epiponini. O BD1 feito das operárias para as rainhas é uma apomorfía das operárias de Metapolybia. Outros comportamentos de teste entre rainhas surgiram independentemente em algumas espécies e poderiam estar relacionados com um aumento na complexidade das interações. Os testes das operárias estariam presentes no ancestral dos Epopinini indicando que elas controlam a seleção das rainhas. O comportamento de worker dance é utilizado principalmente pelas operarias dos clados de Polybia e Metapolybia para testar a capacidade reprodutiva das rainhas. Parece que os comportamentos ritualizados de dominância e de teste são utilizados como sinais honestos do potencial reprodutivo das fêmeas em vez de comportamentos agressivos. Em Metapolybia e Protopolybia exigua o BD1 é realizado com maior frequência após a eliminação da rainha e durante a seleção, diferente de Chartergellus e Protopolybia picteti onde não houve mudança. Em Chartergellus não houve um aumento dos displays, porque as fêmeas no geral não interagiam e a seleção é feita mediante a expulsão das rainhas. Em P. picteti nem todas as rainhas foram eliminadas. Em Metapolybia e Chartergellus no início do ciclo colonial há tolerância reprodutiva, e a eliminação das rainhas estimulou muitas fêmeas a ovipositar, mas o número eventualmente diminui. Este resultado mostra que a delimitação de casta poderia ser feita mediante algum tipo de sinalização química. Em Protopolybia há pouca redução do número de rainhas ao longo do ciclo e as intermediárias constantemente tentam botar ovos. Para as intermediárias é conveniente manter os ovários ativos para tentar atingir o status de rainha em uma nova colônia. A totipotência das fêmeas para desenvolver seus ovários é basal para os Epiponini. As operárias e intermediárias jovens, de idade intermediária e velhas podem desenvolver seus ovários diante da ausência da rainha. A flexibilidade é um traço muito importante nos Epiponini, uma vez que permite à população responder a situações adversas, mobilizando indivíduos a realizar diferentes tarefas segundo as necessidades da população. / Epiponini wasps have colonies with multiple queens, which alternate their number under different stages of the colony cycle. Initially there are several potential queens, but as the colony cycle develops, queen number is reduced. Because most of individuals remain reproductively totipotent there is great potential for conflicts over reproduction. Under this conditions workers could have an advantage to control queen production because they are the dominant caste. Reproductive control could be performed by chemical signals, which along with ritualized behaviors communicate queen presence in the colony. However, queen selection process is a little known across Epiponini, and for this reason our goal was to study queens and workers behavior during queen selection phase, performing a phylogenetic analysis to understand behavioral changes among species. We performed observations in nine species belong to five genera: Chartergellus, Metapolybia, Protopolybia, Polybia, Brachygastra. We marked queens, a sample of workers and newly emerged females to perform direct and video focal observations. To induce queen production, we performed queen removal tests. Some of queens characteristic behaviors and those associated with queen selection process were analyzed via phylogenetic optimization. Bending display I (BD1) is a dominance display performed queens against workers, is the behavior that better identify queens and is also basal for all Epiponini clades. BD1 perform from workers to queens is an apomorphy of Metapolybia. Another test behaviors performed among queens appeared independently in some species, and could be related with an increase of interaction complexity. Worker tests were present in the Epiponini ancestor, indicating that workers are in charge of queen control. Worker dance behavior was synapomorphic for the clade composed by Polybia, Metapolybia and their close related genera, in which workers test queens reproductive potential. Apparently, ritualized tests and dominance behaviors are used in Epiponini as honest signals of queens reproductive potential instead of aggressive behaviors. In Metapolybia and in Protopolybia exigua, BD1 is performed with high frequency after queen elimination and during queen selection process. On the other hand, in Chartergellus and Protopolybia picteti displays frequency did not change. In Chartergellus BD1 frequency did not change because females do not interact to each other a lot, also queen selection is performed by queens expulsion instead by tests. In P. picteti we possibly failed to remove all queens from the colony. Metapolybia and Chartergellus have reproductive tolerance between females during first stages of colony cycle, and queen elimination induced females to lay eggs, but queen number eventually is reduced. These results suggest that reproductive control in fact could be performed by chemical signals. On the other hand, in Protopolybia queens number change little during the colony cycle. In adition, intermediates constantly attempt to lay eggs. Persistence oviposition could be convenient for intermediates in Protopolybia to achieve queen status in new colonies. Females totipotency to develop ovaries is also basal for Epiponini. Not only young females develop their ovaries under queen lost; old and middle age individuals also attempt to replace the queen. Caste flexibility is decisive for colony survival of swarm wasps because allows colonies to respond efficiently to different situations that may eventually arise.

Ropalidia marginata is characterized as a primitively eusocial wasp due to the absence of morphological differentiation between the queen and worker castes. Unlike other primitively eusocial wasps, however, the queen in this species is a docile individual, who does not use aggression to regulate worker reproduction, and does not act as the central pacemaker of her colony. However, if the queen dies or is experimentally removed, one of the workers steps up her aggression immensely within minutes, and if the queen is not replaced, she develops her ovaries, reduces aggression and takes over as the new queen of the colony. We call her the potential queen (PQ). When I started my work on R. marginata, two very intriguing questions were demanding to be answered, which had developed from work done by my immediate seniors in the lab. I decided to pursue both of these for my thesis. My work has been enriched by inputs from several collaborators and colleagues - I couldn’t have done all of it by myself. So, henceforth, I will be using the word “we”, instead of the first person singular to describe the work that has gone into this thesis. Question 1: Is there a designated successor to the queen in R. marginata? My senior Sujata P. Kardile has shown in her thesis, that in R. cyathiformis, a primitively eusocial wasp very closely related to R. marginata, the queen is always succeeded by the next most aggressive individual in the colony, and so the PQ is easily predictable in the presence of the queen. However, in R. marginata, the PQ appears to be an unspecialized individual, who cannot be predicted in the presence of the queen by using age, ovarian profile or behaviour as the yardsticks. However, the PQ becomes evident within minutes after queen removal. The swiftness with which the PQ is established led us to believe that perhaps the successor to the queen in R. marginata is known to the wasps, though we cannot identify her in the presence of the queen. We designed an experiment to check for the presence of such a ”cryptic successor” in R. marginata. Our experiments involved splitting a normal, queen-right nest into two halves separated by a wire mesh partition, so that the wasps could not move across the mesh. Earlier we had used this set-up to demonstrate that a PQ always establishes herself on the queen-less fragment of the nest. So, to test if there is a cryptic successor, we allowed a PQ to establish herself on the queen-less fragment, and then exchanged the queen and the PQ (designated as PQ1) between the two sides. There is a 50% probability that the cryptic successor, if present, would be on the queen-less side in the beginning. Then, upon exchange, she should be able to hold her position on the other side easily. On the other hand, if the cryptic successor is first on the queen-right side, then, upon exchange, she should take over as the PQ (PQ2), and PQ1 should not be able to hold her status. The cryptic successor hypothesis had two predictions: (i) the PQ1 would lose to a PQ2 in about half the cases, and (ii) there would never be a PQ3. We obtained a PQ2 in 5 out of 8 cases, and we never had a PQ3. So we could conclude that there is indeed one individual who is the designated successor to the queen in R. marginata. Since we could not identify her in the presence of the queen, we call her the cryptic successor. The cryptic successor did not receive even a single act of aggression from the PQ1, or from any other individual in the colony. Thus we conclude that she is acceptable to all the wasps in the colony. We next used the more sophisticated and rigorous method of network analysis to check if the PQ could be predicted due to some unique position she might be holding in the social network on her colony. Since this was a first study in a primitively eusocial insect using network tools, we began by characterizing the social networks of queen-right and queen-less colonies of R. marginata, and compared them with the R. cyathiformis networks to see how different the R. marginata society is from a typical primitively eusocial one. The R. marginata social networks based on dominant-subordinate interactions were low in their centrality measure as compared to the R. cyathiformis networks. However, in both the species, the queen-less networks were highly centralized, star-shaped networks with the PQs at the centre. Neither the queens, nor the PQs were key individuals in the queen-right colonies, but it is interesting to note that the removal of an insignificant node, the queen, resulted in a major change in the network architecture, converting the de-centralized queen-right network into a highly centralized one. Such centralized star-shaped networks are unique, and to our knowledge, the first ever described, in any social system. When we removed the queen from the data set (in silico removal), the resulting network was similar in centrality to the queen-right networks. We then did a comparative analysis of the positional importance of the PQs of the two species, and tried to see if we could use this as a tool to predict the PQ in the queen-right network. In R. cyathiformis, the PQs had consistently high ranks (mostly rank 2) in the network based on the degree index, while the PQs in R. marginata had random ranks in the hierarchy. However, since the PQs are known not to have unique ranks in the dominance hierarchies, we repeated the analysis using data on all interactions from the Q-PQ exchange experiments described above. Neither the cryptic successors nor the losers occupied any unique ranks in the all interactions networks. Thus the successors in R. marginata are truly cryptic, even in their social networks. Since R. marginata is known to be more evolved than typical primitively eusocial species, it is likely that the queen’s successor is identified by the wasps through some subtle cue like smell, and so we cannot identify her using the methods that are adequate for the identification of the PQ in a typical primitively eusocial species like R. cyathiformis. Question 2: How does the queen signal her presence and reproductive status to her workers or, how do the workers perceive the presence of their queen? The fact that in spite of her docility, the queen in R. marginata manages to maintain complete reproductive monopoly in her colony, gives rise to the obvious question of how she suppresses worker reproduction. The most attractive hypothesis is that she uses a pheromone like queens of highly eusocial species. My senior A. Sumana had shown that the queen pheromone, if present, is not a volatile substance. She also showed that the queen interacts at a very low rate with her workers, and so they cannot possibly perceive her by means of direct interactions. Since the PQ steps up her aggression within minutes of queen removal, we used her as a proxy to know how soon the queen’s absence is felt in the colony. We built a model to delineate the relationship between the decay time of the pheromone (td), the average age of the queen’s signal present with the PQ (ta), and the average realization time (tr); where tr = td − ta. Using Dijkstra’s algorithm, we showed that the queen could interact faster with the PQ by using relay interactions. Then using experimental data from 50 colonies, we obtained a ta of 102.9 minutes. The td was 340 minutes, and so we obtained a tr of 237.1 minutes; which meant that the PQ should not perceive the queen’s absence within 237 minutes of queen removal, if the queen pheromone is transmitted by a relay mechanism. However, from our experimental data, we had obtained a tr of 30 minutes. So we concluded that physical interactions, both direct and indirect were inadequate for the workers to perceive their queen. As we had ruled out physical interactions, we then wanted to check if it is possible that the queen applies her pheromone to the nest material, from where it is perceived by the workers when they walk or sit on the nest, or antennate the nest surface. The “rub abdomen behaviour (RA)” has been observed to be quite typical of R. marginata queens, and is not very common in the workers of the species. RA involves rubbing the ventral side of the tip of the abdomen or dragging it on the nest surface while walking. We thought that the queen might be using this behaviour to apply her pheromone on the nest material. So we characterized this behaviour using focal behaviour sampling, and found that the queen rubs her abdomen on the nest once in every 23 minutes. Since the observed tr is 30 minutes, it is quite likely that the queen uses the rub abdomen behaviour to apply her pheromone on the nest. The next step was to check for the source of the queen pheromone. We looked for glands that open near the base of the sting, and the Dufour’s gland was a good choice, as it is known to be involved in the recognition of egg-laying workers in the honeybees. We performed a bioassay in the blind using the crude extract of the Dufour’s gland (prepared in Ringer’s solution) from the queen. The Dufour’s gland extract of a randomly chosen worker and the solvent were used as controls. We found that the PQ responds to the queen’s Dufour’s gland extract by lowering her aggression to 65% of what she was showing on queen removal and before the application of the extract. However, the PQ did not change her behaviour significantly when the worker’s extract or Ringer’s solution was applied. The PQ’s reduction of aggression on application of the queen’s extract mimicked the reaction of PQ’s when the queen is re-introduced on the nest some time after removal. So we hypothesize that the Dufour’s gland is the source of the queen pheromone (signal) in R. marginata. This thesis has opened up newer questions pertaining to the power of the queen and the intricacies of the succession to power in R. marginata. For example, we need to pursue chemical analyses of the Dufour’s gland extract of R. marginata to have conclusive proof of it’s being the source of the queen pheromone. But that is perhaps suitable topic for my juniors in the lab, who can continue the tradition of beginning with questions opened up by their seniors!

Submitted by Renata Lopes (renatasil82@gmail.com) on 2017-04-17T18:05:35Z No. of bitstreams: 1 mateusfajardodefreitassalviatodetoni.pdf: 1310680 bytes, checksum: 07d34f3c206819ff6b208c6ab33be11a (MD5) / Approved for entry into archive by Adriana Oliveira (adriana.oliveira@ufjf.edu.br) on 2017-04-18T13:50:54Z (GMT) No. of bitstreams: 1 mateusfajardodefreitassalviatodetoni.pdf: 1310680 bytes, checksum: 07d34f3c206819ff6b208c6ab33be11a (MD5) / Made available in DSpace on 2017-04-18T13:50:54Z (GMT). No. of bitstreams: 1 mateusfajardodefreitassalviatodetoni.pdf: 1310680 bytes, checksum: 07d34f3c206819ff6b208c6ab33be11a (MD5) Previous issue date: 2017 / Os ninhos de insetos sociais funcionam como a interface entre as colônias e o ambiente, e estudar como os fatores ambientais se relacionam com o comportamento de nidificação é essencial para compreender o sucesso desses animais em colonizar e sobreviver na área urbana. Dessa forma, este trabalho teve como objetivo estudar a relação entre a orientação dos ninhos da vespa social Polybia platycephala em áreas urbanas e a incidência de luz ambiental sobre os mesmos, além de descrever sua dinâmica de temperatura em relação ao microambiente em que estão localizados. Para a orientação e a incidência de luz, 11 ninhos foram selecionados em 2016 na cidade de Juiz de Fora, MG, sendo 11 na estação chuvosa e 11 na seca (n= 22). A orientação dos ninhos foi constatada e a incidência de luz de cada ninho foi verificada ao longo do dia (06:00h – 18:00h). Adicionalmente, seis ninhos tiveram sua orientação experimentalmente invertida e a luminosidade foi acompanhada antes e depois da inversão de forma a verificar o efeito da orientação natural sobre a exposição dos ninhos à luz. Para a dinâmica de temperatura utilizaram-se os mesmos 22 ninhos, e a temperatura dos ninhos e do ambiente foram medidas paralelamente à luminosidade. Para verificar o efeito da atividade das vespas sobre a temperatura da colônia, um ninho abandonado e uma colônia ativa foram acompanhados por 24 horas. Os resultados permitiram-nos concluir que P. platycephala apresenta uma orientação de ninhos fortemente enviesada para leste, aumentando a exposição à luz ambiental no período da manhã; são, no entanto, capazes de modificar esse comportamento para adaptar-se ao seu microambiente para otimizar essa exposição. A temperatura dos ninhos é muito relacionada à ambiente, flutuando paralelamente a esta, o que indica uma grande dependência das colônias do mesmo para sua sobrevivência. De forma geral, constatou-se que P. platycephala possui uma íntima relação com o seu microambiente, o que pode ajudar a explicar seu sucesso em colonizar áreas urbanas, mas também a torna sensível a alterações ambientais e climáticas nas mesmas. / Nests of social insects function as the interface between colonies and the environment, and studying how environmental factors relate to the nesting behavior is essential in order to understand these organisms’ success in settling and surviving in the urban area. On this sense, our work aimed to study the relation between the social wasp Polybia platycephala nest orientation and the incidence of environmental light, aside from describing its temperature dynamics regarding the microenvironment where it is set. In order to study nest orientation and light incidence, 22 nests were chosen in 2016 in the city of Juiz de Fora, MG, being 11 in the rainy season and 11 in the dry (n= 22). Nest orientation was verified and light incidence on each nest was assessed through the day (06:00h - 18:00h). Additionally, six nests had their orientation experimentally inverted and luminosity was assessed before and after the inversion in order to verify the effect of nest orientation on exposure to light. In order to study temperature dynamics, the same 22 nests were used, and the nest and ambient temperatures were assessed in parallel with the luminosity. In order to verify the effect of the wasp activity on colony temperatures, an abandoned nest and an active colony were observed for 24 hours. The results allowed us to conclude that P. platycephala shows nest orientation strongly biased towards east, increasing exposure to light during the morning; colonies are, however, able to perform modifications on this behavior in order to adapt themselves to their microenvironment and optimize this exposure. Nest temperatures are intensely related to the ambient, fluctuating in parallel with it, which indicates a great dependence of the colonies on the environment to assure their survival. Overall, we found that P. platycephala has an intimate relation with its microenvironment, which may help explain its success in settling urban areas, but also makes it sensible to environmental and climatic changes in them.

Made available in DSpace on 2014-06-11T19:30:56Z (GMT). No. of bitstreams: 0 Previous issue date: 2009-12-07Bitstream added on 2014-06-13T19:19:44Z : No. of bitstreams: 1 saidemberg_dm_dr_rcla.pdf: 1615145 bytes, checksum: 92ac7814050532bc681c040810a6efc1 (MD5) / Consideráveis esforços estão sendo feitos no sentido de isolar e identificar compostos neuroativos presentes em secreções de Artrópodes, resultando na descoberta de muitos peptídeos e moléculas pequenas com ação bloqueadora de receptores de glutamato e/ou canais de cálcio. Tendo isso em vista, a secreção de muitos desses animais tornaram-se ferramentas úteis para os estudos fisiológicos de diversas funções neurais, e muitos dos compostos neurotóxicos produzidos por suas secreções agressivas/defensivas podem se tornar modelos estruturais e funcionais para o desenvolvimento racional de agentes neuroprotetores para diversas desordens neurológicas. Assim, os venenos das vespas Agelaia pallipes pallipes, Agelaia vicina e Polybia paulista foram fracionados em HPLC, e as frações puras, mais abundantes, foram investigadas por espectrometria de massas e ressonância magnética nuclear para elucidação estrutural das mesmas; e posteriormente, estas frações foram submetidas a diversos ensaios para a determinação de suas atividades biológicas, principalmente daquelas atividades relacionadas à neurotoxicidade/ neuroproteção, tanto em artrópodes como em mamíferos. Através desta abordagem foi possível identificar alguns dos compostos de baixas massas moleculares mais abundantes nos venenos das vespas sociais citadas acima, como a Histamina, a Serotonina, que são comuns a todos estes venenos e possuem como principal função, a potencialização da dor e inflamação, além de outros efeitos no SNC de vertebrados. Também foi isolado um novo tripeptídeo de sequência Gly-Leu-Leu-OH a partir do veneno da vespa A. vicina, cuja função ainda não é conhecida e a síntese e ensaios biológicos do mesmo serão realizados em algumas ramificações dessa tese. A partir do veneno da vespa P. paulista foi isolada 2-feniletilamina, um composto... / Considerable research efforts have been mounted to isolate and identify neuroactive compounds in Arthropods secretions, resulting in the discovery of many peptides and small molecules which block glutamate receptors and/or calcium channels. Thus, the secretion of many of these animals proved to be useful tools for physiological studies of neuronal function, and several of the neurotoxic compounds produced by their defensive/aggressive secretions may become structural and functional models for the rational development of neuroprotective agents for different neurological disorders. Thus, the venoms of Agelaia pallipes pallipes, Agelaia vicina e Polybia paulista were fractionated in a HPLC system, and the most abundant and pure fractions were investigated by Mass Spectrometry and Nuclear Magnetic Resonance for structural elucidation, and then, these compounds were assayed for biological activity determination, focusing the neuronal activities for these compounds, both in Arthropods and mammals. Considering this approach, it was possible to identify some of the most low molecular mass compounds in the venoms of the social wasps cited above, such as: Histamine and Serotonin, which were common to all of these species, presenting as main function to increase the pain and inflammation, in addition to other effects in the CNS of vertebrates. It was also isolated from the venom of the wasp A. vicina a new tripetide which had its sequence determined as Gly-Leu-Leu- OH, and whose function has not been determined. From the venom of the wasp P. paulista it was isolated 2-phenylethylamine, an amphetamine-like compound, whose occurrence is uncommon in animal venoms, and which has stimulatory effects on the CNS primarily related to dopaminergic neurotransmission. It caused effects of paralysis on Africanized bees, probably by hyper-excitability in the CNS of this organism. We have also observed locomotor... (Complete abstract click electronic access below)

Orientador: Mario Sergio Palma / Banca: Emanuel Carrilho / Banca: Dulce Helena Siqueira Silva / Banca: Roberta Cornello Ferreira Nocelli / Banca: Maria Elena de Lima Perez Garcia / Resumo: Consideráveis esforços estão sendo feitos no sentido de isolar e identificar compostos neuroativos presentes em secreções de Artrópodes, resultando na descoberta de muitos peptídeos e moléculas pequenas com ação bloqueadora de receptores de glutamato e/ou canais de cálcio. Tendo isso em vista, a secreção de muitos desses animais tornaram-se ferramentas úteis para os estudos fisiológicos de diversas funções neurais, e muitos dos compostos neurotóxicos produzidos por suas secreções agressivas/defensivas podem se tornar modelos estruturais e funcionais para o desenvolvimento racional de agentes neuroprotetores para diversas desordens neurológicas. Assim, os venenos das vespas Agelaia pallipes pallipes, Agelaia vicina e Polybia paulista foram fracionados em HPLC, e as frações puras, mais abundantes, foram investigadas por espectrometria de massas e ressonância magnética nuclear para elucidação estrutural das mesmas; e posteriormente, estas frações foram submetidas a diversos ensaios para a determinação de suas atividades biológicas, principalmente daquelas atividades relacionadas à neurotoxicidade/ neuroproteção, tanto em artrópodes como em mamíferos. Através desta abordagem foi possível identificar alguns dos compostos de baixas massas moleculares mais abundantes nos venenos das vespas sociais citadas acima, como a Histamina, a Serotonina, que são comuns a todos estes venenos e possuem como principal função, a potencialização da dor e inflamação, além de outros efeitos no SNC de vertebrados. Também foi isolado um novo tripeptídeo de sequência Gly-Leu-Leu-OH a partir do veneno da vespa A. vicina, cuja função ainda não é conhecida e a síntese e ensaios biológicos do mesmo serão realizados em algumas ramificações dessa tese. A partir do veneno da vespa P. paulista foi isolada 2-feniletilamina, um composto... (Resumo completo, clicar acesso eletrônico abaixo) / Abstract: Considerable research efforts have been mounted to isolate and identify neuroactive compounds in Arthropods secretions, resulting in the discovery of many peptides and small molecules which block glutamate receptors and/or calcium channels. Thus, the secretion of many of these animals proved to be useful tools for physiological studies of neuronal function, and several of the neurotoxic compounds produced by their defensive/aggressive secretions may become structural and functional models for the rational development of neuroprotective agents for different neurological disorders. Thus, the venoms of Agelaia pallipes pallipes, Agelaia vicina e Polybia paulista were fractionated in a HPLC system, and the most abundant and pure fractions were investigated by Mass Spectrometry and Nuclear Magnetic Resonance for structural elucidation, and then, these compounds were assayed for biological activity determination, focusing the neuronal activities for these compounds, both in Arthropods and mammals. Considering this approach, it was possible to identify some of the most low molecular mass compounds in the venoms of the social wasps cited above, such as: Histamine and Serotonin, which were common to all of these species, presenting as main function to increase the pain and inflammation, in addition to other effects in the CNS of vertebrates. It was also isolated from the venom of the wasp A. vicina a new tripetide which had its sequence determined as Gly-Leu-Leu- OH, and whose function has not been determined. From the venom of the wasp P. paulista it was isolated 2-phenylethylamine, an amphetamine-like compound, whose occurrence is uncommon in animal venoms, and which has stimulatory effects on the CNS primarily related to dopaminergic neurotransmission. It caused effects of paralysis on Africanized bees, probably by hyper-excitability in the CNS of this organism. We have also observed locomotor... (Complete abstract click electronic access below) / Doutor