Thesis (Ph. D.)--Miami University, Dept. of Botany, 2005. / Title from second page of PDF document. Includes bibliographical references.

京都大学 / 0048 / 新制・課程博士 / 博士(人間・環境学) / 甲第18359号 / 人博第672号 / 新制||人||162(附属図書館) / 25||人博||672(吉田南総合図書館) / 31217 / 京都大学大学院人間・環境学研究科相関環境学専攻 / (主査)教授 市岡 孝朗, 教授 加藤 眞, 教授 松井 正文 / 学位規則第4条第1項該当 / Doctor of Human and Environmental Studies / Kyoto University / DGAM

Coordenação de Aperfeiçoamento de Pessoal de Nível Superior - CAPES / Ecological interactions between organisms are complex and can present dynamic results, which are dependent on environmental context. Plants with extrafloral nectaries attract a wide variety of ants species, in associations commonly considered mutualistic. In tropical environments the combination of plants and ants play an important role in communities. Turnera subulata is a ruderal shrub widely distributed in disturbed areas, which has a pair of extrafloral nectaries in the base of each leaf. In this study, we evaluated whether the ants associated with T. subulata: (i) vary spatially (between locals) and temporally (over day period); (ii) respond to simulate presence of herbivores and the signs of damage caused in different structures of the host plant (stem and leaf); (iii) reduce herbivory rates; (iv) influencing the vegetative growth and reproductive success of the host plant, along its phenology. The experiments were conducted in São Cristóvão-SE. For the initial understanding of the system studied, T. subulata individuals were selected, in which we performed simulations of the presence of herbivorous and realization of injuries in the stem and leaves of the host plants. The total number of individuals and species associated ants was quantified during three periods of the day. It was quantitated the percentage of leaf area lost and leaves attacked by sucking. They also plot experiments were performed in which were established with and without ants. In each plot were transplanted T. subulata seedlings that were followed throughout their phenological stages. Observations were made in order to quantify the vegetative growth and reproductive investment measures the host plant as well as the number of sheets to damage by sucking insects and chewing; and the number of herbivores. The number of visitors associated with T. subulata was quantified during three periods of the day, being held 5min observation / share. In all cases, data were analyzed using general linear models. 21 species of ants were found. The composition of the ant meeting showed significant variation between locations and time of day. Visitation rates and predation by ants was higher in stems than in the leaves of plants. In general, herbivory rates were not correlated with the association / activity of ants, with the exception of leaf area proportion consumed, which showed a significant reduction in plants where the ants defended the leaves. The results showed that the maturation stage there was a trade-off between growth x propagation of plants; which is favorable for plants (ex.: higher reproduction) who were in plots with ants. Plants on treatment with ants had fewer visitors, including predators and parasitoids. The number of sucking herbivores was significantly reduced in the presence of ants, during the flowering stage. Our results suggest that the benefits of the association may be dependent on context. Moreover, they can contribute to the understanding of the mechanisms involved in facultative interactions between ants and plants and to the understanding of the communities in interaction nets. / Interações ecológicas entre organismos são complexas, podendo apresentar resultados dinâmicos, os quais são dependentes do contexto ambiental. Plantas com nectários extraflorais atraem uma ampla variedade de espécies de formigas, em associações comumente consideradas mutualísticas. Nos ambientes tropicais a associação de plantas e formigas desempenha importante papel nas comunidades. Turnera subulata é um arbusto ruderal, amplamente distribuído em áreas antropizadas, que apresenta um par de nectários extraflorais na base de cada folha. Neste estudo, avaliamos se as formigas associadas a T. subulata: (i) variam espacialmente (entre locais) e temporalmente (ao longo do período do dia); (ii) respondem à simulação da presença de herbívoros e aos sinais de danos ocasionados em diferentes estruturas da planta hospedeira (caule e folha); (iii) reduzem as taxas de herbivoria; (iv) influenciam o crescimento vegetativo e sucesso reprodutivo da planta hospedeira, ao longo da sua fenologia. Os experimentos foram realizados em São Cristóvão-SE. Para o entendimento inicial do sistema estudado, foram selecionados indivíduos de T. subulata, nos quais realizamos simulações da presença de herbívoro e realização de injúrias no caule e folhas das plantas hospedeiras. O número total de indivíduos e de espécies de formigas associadas foi quantificado durante três períodos do dia. Foi quantificada a proporção de área foliar perdida e de folhas atacadas por sugador. Também foram realizados experimentos em que foram estabelecidas parcelas com e sem formigas. Em cada parcela foram transplantadas mudas de T. subulata que foram acompanhadas ao longo de seus estágios fenológicos. Foram feitas observações a fim de quantificar as medidas de crescimento vegetativo e investimento reprodutivo da planta hospedeira, assim como o número de folhas com danos por insetos sugadores e mastigadores; e o número de herbívoros. O número de visitantes associados a T. subulata foi quantificado durante três períodos do dia, durante 5min de observação/parcela. Em todos os casos, os dados foram analisados através de modelos lineares generalizados. Foram encontradas 21 espécies de formigas. A composição da assembleia de formigas apresentou variação significativa no espacialmente e temporalmente. As taxas de visitação e de predação pelas formigas foi maior no caule do que nas folhas das plantas. De forma geral, as taxas de herbivoria não foram correlacionadas com a associação/atividade das formigas, com exceção da proporção de área foliar consumida, que mostrou redução significativa em plantas onde as formigas defenderam as folhas. Os resultados mostraram que no estágio de maturação houve um trade-off entre crescimento x reprodução das plantas; sendo este favorável (ex.: maior reprodução) para as plantas que estavam em parcelas com formigas. Plantas sobre tratamento com formigas tiveram menor número de visitantes, incluindo predadores e parasitoides. O número de herbívoros sugadores foi significativamente reduzido na presença de formigas, durante o estágio de floração. Nossos resultados sugerem que os benefícios da associação podem ser dependentes de contexto. Além disso, podem contribuir para a compreensão dos mecanismos envolvidos nas interações facultativas entre formigas e plantas, e para o entendimento das redes de interações em comunidades.

Orientadores: Paulo Sergio Moreira Carvalho de Oliveira, Sebastian Felipe Sendoya Echeverry / Dissertação (mestrado) - Universidade Estadual de Campinas, Instituto de Biologia / Made available in DSpace on 2018-08-23T06:11:49Z (GMT). No. of bitstreams: 1 Vidal_MayraCadorin_M.pdf: 1986442 bytes, checksum: da9ed52c0a73527b0590731f5ee8d10b (MD5) Previous issue date: 2013 / Resumo: Exploradores do mutualismo - indivíduos que utilizam recursos/serviços produzidos pelos mutualistas sem recompensá-lo - podem trazer sérios danos aos mutualistas explorados, principalmente quando acaba matando um dos parceiros mutualistas. Plantas portadoras de NEFs podem manter mutualismos com formigas visitantes, que defendem a planta contra insetos herbívoros. No cerrado de Itirapina (SP), encontramos larvas de uma nova espécie de díptero do gênero Rhinoleucophenga (Drosophilidae) que constroem abrigos de consistência pegajosa em cima dos NEFs de Q. grandiflora. Assim, larvas de Rhinoleucophenga podem interferir no mutualismo formiga-planta, agindo potencialmente como exploradora dessa interação. O presente estudo teve como objetivos principais investigar a história natural dessas larvas, principalmente aspectos do comportamento e interação com Qualea e formigas, e analisar seu possível efeito sobre o mutualismo formiga-Q.grandiflora. Durante observações de campo comprovamos que formigas e outros insetos visitantes dos NEFs podem ficar presos ao abrigo larval e servir de alimento para o díptero. Larvas de Rhinoleucophenga sp. nov. ocorrem em 85% dos indivíduos de Q. grandiflora, principalmente na época chuvosa ocupando preferencialmente nectários ativos, perto do ápice e na face abaxial dos ramos. No levantamento da mirmecofauna visitante de Q. grandiflora encontramos 27 morfoespécies de formigas, sendo as duas mais frequentes Camponotus crassus, e uma espécie do gênero Brachymyrmex, as mesmas que foram mais comumente encontradas presas aos abrigos das larvas mirmecófagas. Vimos que as larvas expõem uma substância líquida na abertura de seu abrigo, que comprovamos possuir composição química muito similar a do néctar extrafloral de Q. grandiflora, o que sugere que as larvas utilizam o néctar da própria planta para atrair suas presas. Na presença de larvas de Rhinoleucophenga, menos formigas visitam as plantas e também por menos tempo. Esse forrageamento diferenciado resultou em menor ataque de formigas a cupins vivos (herbívoros simulados). Além disso, na presença das larvas mirmecófagas houve maior abundância de herbívoros mastigadores e maior área foliar removida por herbívoros. Podemos afirmar que as larvas de Rhinoleucophenga sp. nov. utilizam o recurso da planta sem beneficiá-la. Além disso, as larvas do díptero também prejudicam a planta e suas formigas mutualísticas, uma vez que alimentando-se delas, aumentam a incidência de herbívoros e a herbivoria foliar na planta. Dessa forma, as larvas de Rhinoleucophenga sp. nov. estão agindo como exploradoras e do mutualismo formiga-Qualea grandiflora e predadoras de topo, causando efeito cascata nesse sistema / Abstract: Exploiters of mutualism - individuals that use resources/services offered by mutualists giving nothing in return - can cause serious damages to mutualists, especially when it involves the death of one of the partners. Plants bearing EFNs usually maintain mutualism with aggressive ants, which defend the plant against herbivores. In a cerrado area at Itirapina (SP), we found a new dipteran species of the genus Rhinoleucophenga (Drosophilidae) whose larvae construct sticky shelters on top of active EFNs of Q. grandiflora. Field observations revealed those ants and others insects that visit the EFNs can get trapped at the sticky larval shelters, and are consumed by the larvae. We hypothesized that Rhinoleucophenga larvae could be interfering with the ant-Qualea mutualism, and thus be acting as an exploiter of this interaction. Here, we investigate the natural history of Rhinoleucophenga larvae, mainly its behavior and association with ants and Qualea, and their possible effect on the ant-Qualea mutualism. Larvae of Rhinoleucophenga sp. nov. occur in 85% of the individuals of Qualea grandiflora inspected at Itirapina. Rhinoleucophenga larvae occur mostly during the rainy season, mainly at the apex and abaxial surface of the branches. We found 27 ant species visiting Qualea. The two most frequent visiting species, Brachymyrmex sp. 1 and Camponotus crassus, were most common insects trapped at larval shelters. Chemical analyses revealed that Rhinoleucophenga larvae use Qualea's extrafloral nectar to attract insect prey to their shelters. Qualea branches infested by ant-preying Rhinoleucophenga larvae had ant visitors for less time and in lower numbers than dipteran-free branches. This negative effect on ant foraging activity resulted in decreased levels of ant aggression to live termite-baits (i.e., simulated herbivores) on leaves of dipteran-infested compared to dipteran-free branches. Controlled field experiments demonstrated that branches hosting Rhinoleucophenga larvae had higher numbers of chewing herbivores and higher levels of foliar herbivory than dipteran-free branches. By using Qualea's EFNs as larval shelters and as attractants to ant prey, larvae of Rhinoleucophenga sp. nov. negatively affect both the plant and ant visitors, with cascading effects ultimately resulting in increased herbivore damage to leaves. Thus we can conclude that ant-eating Rhinoleucophenga larvae are acting as exploiters of the mutualism between ants and Q. grandiflora and also as top predator, causing cascade effect on this system / Mestrado / Ecologia / Mestra em Ecologia

Extrafloral nectar (EFN) mediates food-for-protection mutualisms between plants and defensive insects. Senna mexicana var. chapmanii is a perennial legume native to the pine rockland habitats of south Florida. My dissertation focuses on how anthropogenic changes to the pine rocklands might affect EFN production by S. chapmanii, and the outcome of EFN mediated interactions. First, I investigated the influence of time of day, leaf damage, and leaf age on EFN production in S. chapmanii. Plants produced more nectar at night than during the day, and leaf damage resulted in increased EFN production. Furthermore, the response to leaf damage was greater when plants were damaged in the morning than when plants were damaged at night. Damage to young leaves elicited a stronger defensive response than damage to older leaves, in line with optimal defense theory. Second, I conducted a field experiment to determine the effects of ant activity, and light intensity, on herbivory rates, growth, and reproductive fitness in S. chapmanii. In shaded habitats, the presence of ants had no effect on herbivory rates, seed set, or plant size. In sunny habitats, however, plants with ants suffered less herbivore damage, produced more seeds, and grew larger over the duration of the one year study. Third, through a controlled greenhouse experiment I examined the effects of light intensity, and red/far-red light ratios, on EFN production in S. chapmanii. Plants in light-limited conditions produced less EFN, and leaf damage elicited increased EFN production regardless of light conditions. Ratios of red/far-red light, however, did not affect EFN production in either damaged or undamaged plants. Finally, I conducted a field study to determine how ants affect reproductive fitness in S. chapmanii. Over a period of eight months I observed the effects of ants on the activity of herbivores, predators, pollinators, and pre-dispersal seed predators. Relative pollinator efficiency, and rates of pre-dispersal seed predation, were unaffected by ants. Plants with ants, however, were quicker to establish, grew larger, and produced floral displays that attracted more pollinators. In S. chapmanii ants affected plant reproductive fitness simply by facilitating growth and establishment, with coincidental effects on reproductive investment.

Various aspects of myrmecochory were investigated. Effect of different storage methods on atractiveness of seeds with elaiosomes for ants was examined. Viable seed bank of refuse piles (places where ants deposit unused objects from ant nests, e.g. seeds after the elaiosome was eaten) and places outside them were compared. Chemical content of five major chemical groups, specifically amino acids, free fatty acids, organic acids, polyols and sugars, of elaiosomes and seeds of selected species was determined, and compared with respect to their taxonomic relatedness.

Various aspects of myrmecochory were investigated. Effect of different storage methods on atractiveness of seeds with elaiosomes for ants was examined. Viable seed bank of refuse piles (places where ants deposit unused objects from ant nests, e.g. seeds after the elaiosome was eaten) and places outside them were compared. Chemical content of five major chemical groups, specifically amino acids, free fatty acids, organic acids, polyols and sugars, of elaiosomes and seeds of selected species was determined, and compared with respect to their taxonomic relatedness.

Fundação de Amparo a Pesquisa do Estado de Minas Gerais / Ants that collect nectar from plants bearing extrafloral nectaries (EFN) can defend them against the attacks of herbivores. Many cases of this food-for-protection mutualism have been described and the next step is to understand the mechanisms that structure these interactions. In this study, this issue was addressed through field observations and experiments on ant behaviour in studies compiled into five chapters. Specifically, we evaluated the plant\'s investment in attracting ants (nectar quality) and the competitive relationship between ants (resource dominance hierarchies) as structuring mechanisms of ant-plant interactions. The entire study took place in areas of the ecosystem known as Rupestrian Fields, located in Ouro Preto (Brazil), between June 2011 and March 2014. In Chapter 1, we observe that ant-plant interaction network in Rupestrian Fields is nested and generalized as expected for mutualisms, but differs from ant-hemipteran network, its ecological equivalent, despite both share ant species interactions. In Chapter 2, we delve into the study of the effect of resources in interaction frequency, and we showed that the quality more than quantity of nectar determines the pattern of interaction between plants (10 species) and ants (32 species), as well as the benefits of this interaction (conditional outcomes). In the Chapter 3, we focus on the ants to demonstrate, using paired encounters, that ants species are organized in dominance hierarchies based on aggressiveness. This hierarchies affects the numerical dominance based on territoriality and, more importantly, in the dominance of interactions and nectar. Keeping this line of thought, we demonstrated in Chapter 4 that not all ants that collect nectar really protect the plant against herbivores but only the ecological dominant and highly interacted species. Finally, we close the thesis demonstrating in Chapter 5 that the ant species interacted with plants during the day are not the same at night, but this diurnal-nocturnal turnover does not affect the structure of the interactions by only changing the dominant species. This study brings behavioural and empirical evidences on the importance of the identity of the ant species and its ecological dominance, as well the effect of the nectar quality, for the structure of ant-plant interactions and effectiveness of mutualisms. / As formigas que recolhem o néctar das plantas com nectários extraflorais (EFN) podem defendê-las contra ataques de herbívoros. Muitos casos desse tipo de mutualismo foram descritos mas ainda é preciso entender os mecanismos que estruturam essas interações. Neste estudo, esse problema foi abordado através de observações de campo e experimentos sobre o comportamento das formigas compilados em cinco capítulos. Especificamente, foi avaliado o investimento da planta em atrair formigas (qualidade do néctar) e a relação competitiva entre formigas pelo néctar (hierarquias de dominância) como mecanismos estruturadores das interações formiga-planta (partição das interações). Todo o estudo ocorreu em áreas do ecossistema Campo Rupestre, localizadas em Ouro Preto (Brasil), de junho de 2011 a março de 2014. No Capítulo 1, observamos que redes de interação formiga-planta em Campo rupestre é aninhada e generalizada como esperado para mutualismos, mas difere da rede formiga-hemíptero, seu equivalente ecológico, apesar de compartilharem as mesmas espécies de formigas interagentes. No Capítulo 2, nós aprofundamos no efeito dos recursos estruturando interações, e mostramos que a qualidade mais do que a quantidade de néctar determina o padrão de interação entre formigas (32 espécies) e plantas (10 espécies), bem como os benefícios desta interação (benefícios condicionais). No Capítulo 3, nós demonstramos através de encontros interespecíficos pareados que as formigas são organizadas em hierarquias de dominância comportamental baseadas em agressividade. Essa hierarquia afeta a dominância numérica por territorialidade e, mais importante, o domínio das interações e do néctar. Mantendo esta linha de pensamento, demonstramos no Capítulo 4 que nem todas as formigas que coletam néctar realmente protegem a planta contra herbívoros mas somente as espécies dominantes e altamente interagidas. Finalmente, fechamos a tese demonstrando no Capítulo 5 que as espécies de formiga que interagem com as plantas durante o dia não são o mesmo à noite, mas esta rotatividade diurno-noturno não afeta a estrutura das interações devido a troca específica da formiga dominante. Este estudo traz evidências comportamentais e empíricas sobre a importância da identidade de espécies de formiga e sua dominância ecológica, assim como da qualidade do recurso, para a estrutura de rede da formiga-planta e eficácia do mutualismo. / Doutor em Ecologia e Conservação de Recursos Naturais