The sacoglossan sea slug Elysia clarki is able to photosynthesize for three to four months using chloroplasts sequestered from its algal food sources. Furthermore, the slug is able to store multiple chloroplasts from different algal species within the same cell. This research, consisting of several related studies, explores the role that provision of organic nutrients via photosynthesis plays in the biology of the slug. The first chapter demonstrates that, under conditions of starvation, photosynthetic activity in E. clarki remains fully functional for one month after which it then declines. During the first month of starvation the slug exhibits similar feeding behavior as slugs provided a continuous supply of food, suggesting that photosynthesis delays the onset of starvation-induced behavioral changes. The second chapter explores E. clarki's spatial relationships with algae known to be food sources in the field. In areas with high slug density, edible algal populations were very low. DNA barcoding was employed to demonstrate that the algae found near slugs were poor predictors of which foods were actually consumed by slugs. Generally, there was a mismatch between algae available in the field and slug diets. The third chapter explores how E. clarki is able to maintain photosynthesis. After labeling with a C14 ALA incubation process, then chlorophyll was extracted from slugs and purified using HPLC. Results indicate that recently collected E. clarki are able to synthesize chlorophyll, whereas slugs starved for 3 months were not. Photosynthesis plays a very important role for E. clarki and its relationships with food algae.

Este trabalho focou nos aspectos evolutivos e ecológicos dos tricomas em espécies da tribo Bignonieae (Bignoniaceae), e no potencial papel defensivo dessas estruturas no aumento da resistência das plantas frente aos herbívoros. Esta tese caracterizou quatro morfotipos de tricomas encontrados nas partes vegetativas das plantas da tribo Bignonieae morfologicamente, estudou os padrões macro-evolutivos destes tricomas e o papel funcional dos tricomas pateliformes secretores de néctar em diferentes escalas de tempo e espaço. O primeiro capítulo descreveu os quatro tipos de tricomas micro e macro-morfologicamente, e sua relação com a história filogenética das espécies da tribo Bignonieae. Quatro tipos de tricomas distintos foram reconhecidos: tricomas não glandulares (ng), tricomas glandulares peltados (gp), tricomas glandulares estipitados (ge), e tricomas glandulares pateliformes/cupulares (P/C). Três destes tricomas provavelmente já estavam presentes no ancestral da tribo Bignonieae (i.e., não glandulares, glandulares peltados e glandulares pateliformes/cupulares) enquanto os tricomas glandulares estipitados surgiram mais recentemente e múltiplas vezes dentro da tribo. Além disso, realizamos uma revisão das terminologias aplicadas a esses tricomas bem como sugerimos uma padronização dos tipos de tricomas para o grupo de forma a diminuir a divergência entre os trabalhos já publicados. O segundo capítulo testou a eficiência dos tricomas pateliformes secretores de néctar (nectários extraflorais) em duas espécies de Anemopaegma das savanas brasileiras. Essas duas espécies atraíram mais formigas que plantas vizinhas, com os indivíduos de Anemopaegma com maiores quantidades de nectários sendo visitados por um número maior de formigas que as plantas com um número menor de nectários (variação intra-populacional). No entanto, não foi possível observar um efeito dos nectários e formigas sobre a herbivoria e o desempenho das plantas como esperado pela hipótese de defesa mediada por essas estruturas. Hipóteses alternativas foram utilizadas para discutir os resultados, incluindo uma discussão sobre o possível custo/benefício de tais estruturas em diferentes ambientes (florestas e savanas) relacionados com a história filogenética do grupo, além da variação das interações esperada entre populações segundo a hipótese de mosaico geográfico. O terceiro capítulo testou o papel defensivo do sistema planta-formiga mediado pelos nectários extraflorais no contexto filogenético da tribo Bignonieae. Espécies de plantas com um número maior de nectários foram visitadas por um número maior de formigas (teste controlado pelas relações de parentesco entre as espécies). Além disso, espécies mais proximamente relacionadas apresentaram uma diferença na abundância de nectários extraflorais maior do que o esperado pelo modelo neutro de evolução, gerando um padrão de convergência dessas estruturas na tribo Bignonieae. Esse desvio no padrão evolutivo esperado pode ter sido causado por forças direcionais de seleção e momentos de contra-seleção, dado o balanço entre custo-benefício dos nectários extraflorais para as plantas. Neste contexto, dois fatores foram testados para explicar o desvio na evolução da abundância de nectários na tribo Bignonieae: (1) mudança de habitat das florestas para as savannas (fatores extrínsecos); (2) surgimento de novos caracteres morfológicos como outros tipos de tricomas na superfície das plantas (fatores intrínsecos). Ambos fatores podem ter interferido nas interações formiga-planta e no padrão de evolução dos nectários. Dessa forma, a ocupação das savanas levou a uma diminuição do número de nectários (provável contra-seleção dessas estruturas), enquanto o surgimento de tricomas glandulares adesivos teve o mesmo efeito sobre os nectários. Ambos resultados são discutidos considerando a condicionalidade das interações em função da variação biótica (formigas e herbívoros) entre habitats, e também do \"trade-off\" entre os caracteres de defesa. O quarto capítulo testou a teoria de coevolução em mosaico geográfico no sistema planta-formiga-herbívoro em 10 populações da espécie de savana Anemopaegma álbum. Não foram encontradas correlações entre os nectários (e variáveis descritoras do néctar), a abundância de formigas visitantes, a herbivoria ou as variáveis de performance das plantas entre as populações. Esse padrão esteve associado principalmente a variação na assembléia de formigas, a qual foi dominada por formigas do gênero Crematogaster em uma das populações, mas dominadas por formigas do gênero Camponotus na grande maioria das outras populações. No entanto, 3 das 10 populações estudadas apresentaram um alto número de plantas sem formigas, diminuindo muito as chances dessas populações serem defendidas frente aos herbívoros pelos nectários. A abundância de formigas esteve relacionada negativamente com a herbivoria, e positivamente com as variáveis de performance das plantas entre as populações. Das 10 populações amostradas, 5 delas tiveram os nectários acoplados (do inglês, \"matched\") com as formigas visitantes, embora o tipo de acoplamento tenha variado entre elas. Dessas 5 populações, somente 3 tiveram uma produção positiva de folhas e baixa herbivoria, no qual duas delas tiveram alta abundância de nectários nas folhas e foram dominadas por formigas do gênero Camponotus (maiores em tamanho, mas com baixa capacidade de recrutamento). Já a terceira população teve em média a menor abundância de nectários nas folhas, e a mesma apresentou o maior número de formigas por planta (e maior freqüência), em geral formigas do gênero Crematogaster (menores em tamanho mas com grande capacidade de recrutamento). Por serem menores em tamanho, tais formigas utilizaram a secreção dos nectários quase que individualmente. Nem a abundância de nectários, nem as formigas ou mesmo a herbivoria estiveram estruturados espacialmente, corroborando, a hipótese de mosaico geográfico para as interações formiga-planta-herbívoro em A.album. Neste contexto, as três populações com as interações formiga-planta \"mached\" foram consideradas \"hot-spots\" das interações, nas quais as populações atingiram os valores mais altos das variáveis de performance das plantas, enquanto as demais foram consideradas \"cold-spots\". A maioria das populações \"cold-spots\" foi explicada pela falta de formigas suficientes para que as interações com as plantas pudessem se tornar efetivas na defesa frente aos herbívoros, mas outros processos também foram considerados para discutir os resultados apresentados neste trabalho. / This thesis focused on the evolutionary-ecology of trichomes in the tribe Bignonieae (Bignoniaceae), and in the potential defensive role of these structures against herbivores. More specifically, we characterized four trichome morphotypes found in vegetative plant parts of representatives of the Bignonieae, as well as studied the macro-evolutionary patterns of these trichomes, and the functional role of patelliforme nectar secreting trichomes in different scales of time and space. The first chapter describes four different trichome types micro and macromorphologically, as well as investigates their evolutionary patterns during the history of Bignonieae. The four different trichome types recognized are: non-glandular trichomes (ng), glandular peltate trichomes (gp), glandular stipitate trichomes (gst), and glandular patteliform/cupular trichomes (P/Cgt). Our analyses indicated that three of these trichomes were likely already present in the most recent common ancestor of the tribe Bignonieae (i.e., non glandular, glandular peltate, and glandular patteliform/cupular), while the glandular stipitate trichomes evolved more recently and multiple times during the history of the tribe. Results from this study were combined with a literature review in order to revise the trichome terminology and propose standardized names for the various trichome types currently found in the group. The second chapter tested the efficiency of patelliform nectar-secreting trichomes (extrafloral nectaries, EFNs) in two species of Anemopaegma of the Brazilian savannas. These two species attracted more ants than neighboring plants, with individuals that presented higher amounts of EFNs being visited by a higher number of ants than plants with lower amounts of EFNs (intra-population variation). Nonetheless, no effect of EFNs and ants was observed on herbivory nor on the performance of the studied plants, contradicting the expectations of the mediated EFNs defense hypothesis. Alternative hypotheses were also considered including the cost/benefits model to understand the outcomes of ant-plant interaction: (1) phylogenetic inertia hypothesis that connect EFNs-ant interactions with the plant transitions between different environments (forests to savannas); and (2) geographic mosaic hypothesis that predict differences in the outcomes of ant-plant interactions across populations. The third chapter tested the defensive role of extrafloral nectaries in the context of the phylogenetic history of Bignonieae. Species of plants with a higher number of EFNs were visited by a higher number of ants (test controlled by phylogeny). In addition, closely related species presented a higher difference in the abundance of EFNs than expected under the neutral model of evolution. Such deviation may have resulted by directional forces of slection and moments of counter-selection, given the costs and benefits of the extrafloral nectaries for the plants. Two specific factors were considered as the major possible determinants of the evolutionary patterns of the EFNs: (1) change of habitat from forests to savannas (extrinsic factors); and (2) emergence of new morphological characters such as other trichome types over the plant\'s surface (intrinsic factors). Both factors might have altered the ant-plant interactions and the evolution of nectaries. The occupation of the savannas was associated with a decrease in the number of nectaries (likely due to counter-selection of these structures), while the evolution of adhesive glandular trichomes presenting the same effect on the nectaries. Both results are discussed in the light of the biotic variation (ants and herbivores) encountered between habitats, as well as in the light of the trade-off among defensive characters. The fourth chapter tested the geographic mosaic theory of coevolution in 10 populations of the savanna species Anemopaegma álbum. No correlations were found among extrafloral nectaries (and nectar variables), the abundance of visiting ants, herbivory, and plant performance among populations. This pattern was mainly associated with the variation in the assembly of ants encountered in the various populations. Most populations of A. álbum were dominated by assemblages of Camponotus ants, except for one that was dominated by Crematogaster ants. However, 3 of 10 populations studied presented a high number of plants without ants, decreasing the chances of defense against herbivores by EFNs. The abundance of ants was negatively associated with herbivory, and positively associated with plant performance variables among populations. Out of the 10 populations sampled, five presented an abundance of EFNs that matched the functional traits of ants. Out of these five populations, only three presented positive leaf production and low herbivory. From these three populations, two presented high abundances of EFNs on the leaves and were dominated by Camponotus ants (i.e., bigger in size, but with a low recruiting capacity). The third population presented on average the smallest abundance of EFNs on the leaves; it also presented the highest number of ants per plant (and the highest frequency), which generally were Crematogaster ants (smaller in size but with greater recruiting capacity). Because these ants are smaller in size, they used nearly all isolated EFNs encountered over the plant\'s surface. Neither the abundance of EFNs, the abundance of ants, and the assemblage of herbivores were structured spatially, corroborating the geographic mosaic hypothesis for the ant-plant-herbivore interactions in A. album. In this context, the three populations with \"matched\" ant-plant interactions were considered \'hot-spots\' of interactions, in which the populations reached the highest values of plant performance, while the others were considered \'cold-spots\'. The majority of \'cold spot\' populations were explained by a lack of sufficient ants to protect the plants effectively

<p>Plant-animal interactions are affected by both abundance and distribution of interacting species and the community context in which they occur. However, the relative importance of these factors is poorly known. I examined the effects of predator host range, environmental factors, host plant populations, plant traits and fruit abortion on the intensity of pre-dispersal seed predation in 46 host populations of the perennial herb <i>Lathyrus vernus</i>. I recorded damage by beetle pre-dispersal seed predators, mainly <i>Apion opeticum </i>and <i>Bruchus atomarius</i> with different host ranges on <i>L. vernus</i> as well as on two additional host plants. Local seed predator population size was mainly influenced by plant population size, current seed production and beetle population size in the previous year, but was not strongly affected by connectivity. The monophagous seed predator was less abundant and had lower densities than the oligophagous. Both predator species had a strong ability to track fluctuations in seed production; intensity of predation increased with relative increases in seed production. Oligophagous predation on<i> L. vernus i</i>ncreased with the abundance of alternative hosts, but presence of<i> L. vernus</i> did not affect predation on alternative hosts. Abundances and trait preferences differed among three co-occurring seed predators, but were also associated with the abundance of the other species. Overall, seed predation influenced selection on flower number. I found clear indications of seed predator offence but no obvious plant defence. The pattern of fruit abortion was associated with reduced plant fitness since the seed predator had an advanced ability to locate fruits with high probability of retention. Taken together, different factors influencing abundance of the seed predator species, different preferences, and context dependent trait selection are likely to result in complex spatio-temporal variation in overall seed losses and trait selection in the common host plant.</p>

<p>In this thesis, I combined comparative and experimental approaches to examine selection on reproductive traits, and population differentiation in the insect-pollinated, outcrossing, perennial herb <i>Arabidopsis lyrata</i>. More specifically, I (1) determined whether selection on flowering phenology and floral display can be attributed to interactions with pollinators and herbivores, (2) examined whether population differentiation in flowering phenology and floral display is correlated with current selection on these traits, and (3) tested for local adaptation from contrasting environments in Europe.</p><p>A field experiment conducted in a Swedish population demonstrated, that interactions with pollinators may markedly affect selection on both floral display and phenology of flowering. In an alpine population in Norway, grazing damage to inflorescences strongly influenced selection on floral display. The results suggest that variation in the abundance of pollinators and herbivores should contribute to spatio-temporal variation in selection on flowering phenology and floral display in <i>A. lyrata</i>. </p><p>A common-garden experiment showed that flowering phenology and floral display vary among Scandinavian populations of <i>A. lyrata</i>. For some traits patterns of population differentiation were consistent with differences in the direction and strength of phenotypic selection determined in comparisons (a) between an alpine population in Norway and a coastal population in Sweden, and (b) among coastal populations in Sweden. This suggests that current selection contributes to the maintenance of genetic differentiation in these traits.</p><p>Adaptive differentiation among populations was examined in a reciprocal transplant experiment that included populations from three contrasting environments, alpine Norway, coastal Sweden and lowland, continental Germany. The experiment provided evidence for local adaptation, and indicated that populations have diverged in traits affecting plant establishment and early growth.</p>

In this thesis, I combined comparative and experimental approaches to examine selection on reproductive traits, and population differentiation in the insect-pollinated, outcrossing, perennial herb Arabidopsis lyrata. More specifically, I (1) determined whether selection on flowering phenology and floral display can be attributed to interactions with pollinators and herbivores, (2) examined whether population differentiation in flowering phenology and floral display is correlated with current selection on these traits, and (3) tested for local adaptation from contrasting environments in Europe. A field experiment conducted in a Swedish population demonstrated, that interactions with pollinators may markedly affect selection on both floral display and phenology of flowering. In an alpine population in Norway, grazing damage to inflorescences strongly influenced selection on floral display. The results suggest that variation in the abundance of pollinators and herbivores should contribute to spatio-temporal variation in selection on flowering phenology and floral display in A. lyrata. A common-garden experiment showed that flowering phenology and floral display vary among Scandinavian populations of A. lyrata. For some traits patterns of population differentiation were consistent with differences in the direction and strength of phenotypic selection determined in comparisons (a) between an alpine population in Norway and a coastal population in Sweden, and (b) among coastal populations in Sweden. This suggests that current selection contributes to the maintenance of genetic differentiation in these traits. Adaptive differentiation among populations was examined in a reciprocal transplant experiment that included populations from three contrasting environments, alpine Norway, coastal Sweden and lowland, continental Germany. The experiment provided evidence for local adaptation, and indicated that populations have diverged in traits affecting plant establishment and early growth.

Plant-animal interactions are affected by both abundance and distribution of interacting species and the community context in which they occur. However, the relative importance of these factors is poorly known. I examined the effects of predator host range, environmental factors, host plant populations, plant traits and fruit abortion on the intensity of pre-dispersal seed predation in 46 host populations of the perennial herb Lathyrus vernus. I recorded damage by beetle pre-dispersal seed predators, mainly Apion opeticum and Bruchus atomarius with different host ranges on L. vernus as well as on two additional host plants. Local seed predator population size was mainly influenced by plant population size, current seed production and beetle population size in the previous year, but was not strongly affected by connectivity. The monophagous seed predator was less abundant and had lower densities than the oligophagous. Both predator species had a strong ability to track fluctuations in seed production; intensity of predation increased with relative increases in seed production. Oligophagous predation on L. vernus increased with the abundance of alternative hosts, but presence of L. vernus did not affect predation on alternative hosts. Abundances and trait preferences differed among three co-occurring seed predators, but were also associated with the abundance of the other species. Overall, seed predation influenced selection on flower number. I found clear indications of seed predator offence but no obvious plant defence. The pattern of fruit abortion was associated with reduced plant fitness since the seed predator had an advanced ability to locate fruits with high probability of retention. Taken together, different factors influencing abundance of the seed predator species, different preferences, and context dependent trait selection are likely to result in complex spatio-temporal variation in overall seed losses and trait selection in the common host plant.

In this thesis, I examine variation in occurrence and performance of glabrous and trichome-producing plants of the perennial herb Arabidopsis lyrata. I combine field studies and genetic analysis to (1) examine the function of trichomes as a resistance character in natural populations, (2) compare the magnitude of population differentiation in trichome-production and at putatively neutral marker loci, (3) examine the molecular genetic basis of trichome-production, and (4) quantify the effects of herbivore removal on population growth and relative performance of glabrous and trichome-producing plants. In a survey of 30 populations of A. lyrata in Norway and Sweden, I documented spatiotemporal variation in damage from insect herbivores. With few exceptions, glabrous plants were more damaged by herbivorous insects than trichome-producing plants in polymorphic populations. Damage levels varied substantially among populations and among years. The intensity of herbivory quantified as mean leaf removal to glabrous plants was higher in polymorphic populations than in monomorphic glabrous populations. Within the Swedish range, populations were more strongly differentiated at the locus coding for glabrousness than at eight putatively neutral isozyme loci. This is consistent with the hypothesis that trichome production is subject to divergent selection. A study of the genetic basis of trichome production showed that glabrousness was associated with mutations in an orthologue to GLABROUS1, a regulatory gene known to cause glabrousness in A. thaliana. Comparative data indicate that the genetic basis of glabrousness varies among populations. Experimental removal of insect herbivores in a natural A. lyrata population increased population growth rate and the relative fitness of the glabrous morph. The results suggest that insect herbivory may influence both population dynamics and selection on trichome production in A. lyrata.

The strength and direction of interactions between organisms vary spatially across the landscape. Traditionally, the focus has been on how trait variation affects the interactions between species. However, differences in abiotic and biotic environmental factors may also alter the distribution, phenology and behavior of the interacting species. To be able to understand why an interaction varies across the landscape, the effects of trait variation has to be separated from the effects of the environmental context. In this thesis, I try to separate the effects of context and trait differences on plant resistance against herbivory, through experimental and observational studies conducted with two cytotypes of the perennial herb Cardamine pratensis and its main herbivore, Anthocharis cardamines. The results show that differences in plant resistance against oviposition under controlled conditions were mainly mediated by flower size; larger flowers were more attractive to the female butterfly. However, among-populations differences in oviposition under natural conditions were not related to the resistance observed under controlled conditions, or to ploidy type, flowering phenology or plant size. Within populations under natural conditions the oviposition patterns by A. cardamines was affected by the plant traits plant size and flowering phenology. The result of this thesis shows that among-population differences in intensity of plant-herbivore interactions were caused by differences in environmental context rather than by herbivore preferences for any phenotypic plant traits, while host plant selection within population was based on plant traits. This suggests that biotic and biotic context can have important effects on the intensity of plant-herbivore interactions. Although genetic traits influenced the outcome of the interaction within populations, it was the environmental context of the populations that determined largely if the interaction took place or not. / <p>At the time of the doctoral defence the following papers were unpublished and had  a status as follows: Paper 2: Manuscript in review in Plos One; Paper 3: Manuscript in review in <em></em>Ecological Entomology; Paper 4: Manuscript</p>

The dispersal services of frugivores affect plant community assembly, persistence, and gene flow in the short-term, and in the long-term are critical to ensuring that tropical trees and palms can regenerate in disturbed areas and can migrate amidst climate change. Halmahera is the largest Moluccan island within the Wallacea biodiversity hotspot, yet data on its plant and animal distributions and interactions are almost null. I studied the tropical trees and palms of Halmahera and their seed dispersal dynamics. Chapter I explores the palms of the Moluccan islands through field-, herbarium-, and literature- based studies. The results of herbarium specimen collections are presented within a preliminary list of palms for Halmahera’s Aketajawe-Lolobata National Park, and contextualized in a review of regional palm biogeography. Expanding beyond the study of one plant family, Chapter II compiles and examines all the tree and palm taxa of the Moluccan islands in order to infer seed dispersal syndromes for each taxon, resulting in an analysis of over 900 taxa. Zoochory was found in 93% of plant families, and nearly 30% of endemic taxa rely primarily on dispersal by large-bodied frugivores. The role of a hypothesized keystone disperser (the Papuan hornbill, Rhyticeros plicatus ruficollis) is confirmed experimentally to disperse about 10% of Halmahera’s tropical tree and palm taxa. The final chapter encompasses an ex-situ germination trial and a year-long in-situ experimental study that examines the germination and recruitment of three palm species post-ingestion by Papuan hornbills, in four different habitat types representative of Halmahera’s landscape. Palms dispersed by hornbills into disturbed habitats and primary forests resulted in enhanced recruitment, although results varied by habitat and species. Hornbills aid both in forest conservation and recovery/assembly after disturbance. The results of this dissertation provide a foundation for further ecological studies and for enhanced conservation of Halmahera island.

Este trabalho focou nos aspectos evolutivos e ecológicos dos tricomas em espécies da tribo Bignonieae (Bignoniaceae), e no potencial papel defensivo dessas estruturas no aumento da resistência das plantas frente aos herbívoros. Esta tese caracterizou quatro morfotipos de tricomas encontrados nas partes vegetativas das plantas da tribo Bignonieae morfologicamente, estudou os padrões macro-evolutivos destes tricomas e o papel funcional dos tricomas pateliformes secretores de néctar em diferentes escalas de tempo e espaço. O primeiro capítulo descreveu os quatro tipos de tricomas micro e macro-morfologicamente, e sua relação com a história filogenética das espécies da tribo Bignonieae. Quatro tipos de tricomas distintos foram reconhecidos: tricomas não glandulares (ng), tricomas glandulares peltados (gp), tricomas glandulares estipitados (ge), e tricomas glandulares pateliformes/cupulares (P/C). Três destes tricomas provavelmente já estavam presentes no ancestral da tribo Bignonieae (i.e., não glandulares, glandulares peltados e glandulares pateliformes/cupulares) enquanto os tricomas glandulares estipitados surgiram mais recentemente e múltiplas vezes dentro da tribo. Além disso, realizamos uma revisão das terminologias aplicadas a esses tricomas bem como sugerimos uma padronização dos tipos de tricomas para o grupo de forma a diminuir a divergência entre os trabalhos já publicados. O segundo capítulo testou a eficiência dos tricomas pateliformes secretores de néctar (nectários extraflorais) em duas espécies de Anemopaegma das savanas brasileiras. Essas duas espécies atraíram mais formigas que plantas vizinhas, com os indivíduos de Anemopaegma com maiores quantidades de nectários sendo visitados por um número maior de formigas que as plantas com um número menor de nectários (variação intra-populacional). No entanto, não foi possível observar um efeito dos nectários e formigas sobre a herbivoria e o desempenho das plantas como esperado pela hipótese de defesa mediada por essas estruturas. Hipóteses alternativas foram utilizadas para discutir os resultados, incluindo uma discussão sobre o possível custo/benefício de tais estruturas em diferentes ambientes (florestas e savanas) relacionados com a história filogenética do grupo, além da variação das interações esperada entre populações segundo a hipótese de mosaico geográfico. O terceiro capítulo testou o papel defensivo do sistema planta-formiga mediado pelos nectários extraflorais no contexto filogenético da tribo Bignonieae. Espécies de plantas com um número maior de nectários foram visitadas por um número maior de formigas (teste controlado pelas relações de parentesco entre as espécies). Além disso, espécies mais proximamente relacionadas apresentaram uma diferença na abundância de nectários extraflorais maior do que o esperado pelo modelo neutro de evolução, gerando um padrão de convergência dessas estruturas na tribo Bignonieae. Esse desvio no padrão evolutivo esperado pode ter sido causado por forças direcionais de seleção e momentos de contra-seleção, dado o balanço entre custo-benefício dos nectários extraflorais para as plantas. Neste contexto, dois fatores foram testados para explicar o desvio na evolução da abundância de nectários na tribo Bignonieae: (1) mudança de habitat das florestas para as savannas (fatores extrínsecos); (2) surgimento de novos caracteres morfológicos como outros tipos de tricomas na superfície das plantas (fatores intrínsecos). Ambos fatores podem ter interferido nas interações formiga-planta e no padrão de evolução dos nectários. Dessa forma, a ocupação das savanas levou a uma diminuição do número de nectários (provável contra-seleção dessas estruturas), enquanto o surgimento de tricomas glandulares adesivos teve o mesmo efeito sobre os nectários. Ambos resultados são discutidos considerando a condicionalidade das interações em função da variação biótica (formigas e herbívoros) entre habitats, e também do \"trade-off\" entre os caracteres de defesa. O quarto capítulo testou a teoria de coevolução em mosaico geográfico no sistema planta-formiga-herbívoro em 10 populações da espécie de savana Anemopaegma álbum. Não foram encontradas correlações entre os nectários (e variáveis descritoras do néctar), a abundância de formigas visitantes, a herbivoria ou as variáveis de performance das plantas entre as populações. Esse padrão esteve associado principalmente a variação na assembléia de formigas, a qual foi dominada por formigas do gênero Crematogaster em uma das populações, mas dominadas por formigas do gênero Camponotus na grande maioria das outras populações. No entanto, 3 das 10 populações estudadas apresentaram um alto número de plantas sem formigas, diminuindo muito as chances dessas populações serem defendidas frente aos herbívoros pelos nectários. A abundância de formigas esteve relacionada negativamente com a herbivoria, e positivamente com as variáveis de performance das plantas entre as populações. Das 10 populações amostradas, 5 delas tiveram os nectários acoplados (do inglês, \"matched\") com as formigas visitantes, embora o tipo de acoplamento tenha variado entre elas. Dessas 5 populações, somente 3 tiveram uma produção positiva de folhas e baixa herbivoria, no qual duas delas tiveram alta abundância de nectários nas folhas e foram dominadas por formigas do gênero Camponotus (maiores em tamanho, mas com baixa capacidade de recrutamento). Já a terceira população teve em média a menor abundância de nectários nas folhas, e a mesma apresentou o maior número de formigas por planta (e maior freqüência), em geral formigas do gênero Crematogaster (menores em tamanho mas com grande capacidade de recrutamento). Por serem menores em tamanho, tais formigas utilizaram a secreção dos nectários quase que individualmente. Nem a abundância de nectários, nem as formigas ou mesmo a herbivoria estiveram estruturados espacialmente, corroborando, a hipótese de mosaico geográfico para as interações formiga-planta-herbívoro em A.album. Neste contexto, as três populações com as interações formiga-planta \"mached\" foram consideradas \"hot-spots\" das interações, nas quais as populações atingiram os valores mais altos das variáveis de performance das plantas, enquanto as demais foram consideradas \"cold-spots\". A maioria das populações \"cold-spots\" foi explicada pela falta de formigas suficientes para que as interações com as plantas pudessem se tornar efetivas na defesa frente aos herbívoros, mas outros processos também foram considerados para discutir os resultados apresentados neste trabalho. / This thesis focused on the evolutionary-ecology of trichomes in the tribe Bignonieae (Bignoniaceae), and in the potential defensive role of these structures against herbivores. More specifically, we characterized four trichome morphotypes found in vegetative plant parts of representatives of the Bignonieae, as well as studied the macro-evolutionary patterns of these trichomes, and the functional role of patelliforme nectar secreting trichomes in different scales of time and space. The first chapter describes four different trichome types micro and macromorphologically, as well as investigates their evolutionary patterns during the history of Bignonieae. The four different trichome types recognized are: non-glandular trichomes (ng), glandular peltate trichomes (gp), glandular stipitate trichomes (gst), and glandular patteliform/cupular trichomes (P/Cgt). Our analyses indicated that three of these trichomes were likely already present in the most recent common ancestor of the tribe Bignonieae (i.e., non glandular, glandular peltate, and glandular patteliform/cupular), while the glandular stipitate trichomes evolved more recently and multiple times during the history of the tribe. Results from this study were combined with a literature review in order to revise the trichome terminology and propose standardized names for the various trichome types currently found in the group. The second chapter tested the efficiency of patelliform nectar-secreting trichomes (extrafloral nectaries, EFNs) in two species of Anemopaegma of the Brazilian savannas. These two species attracted more ants than neighboring plants, with individuals that presented higher amounts of EFNs being visited by a higher number of ants than plants with lower amounts of EFNs (intra-population variation). Nonetheless, no effect of EFNs and ants was observed on herbivory nor on the performance of the studied plants, contradicting the expectations of the mediated EFNs defense hypothesis. Alternative hypotheses were also considered including the cost/benefits model to understand the outcomes of ant-plant interaction: (1) phylogenetic inertia hypothesis that connect EFNs-ant interactions with the plant transitions between different environments (forests to savannas); and (2) geographic mosaic hypothesis that predict differences in the outcomes of ant-plant interactions across populations. The third chapter tested the defensive role of extrafloral nectaries in the context of the phylogenetic history of Bignonieae. Species of plants with a higher number of EFNs were visited by a higher number of ants (test controlled by phylogeny). In addition, closely related species presented a higher difference in the abundance of EFNs than expected under the neutral model of evolution. Such deviation may have resulted by directional forces of slection and moments of counter-selection, given the costs and benefits of the extrafloral nectaries for the plants. Two specific factors were considered as the major possible determinants of the evolutionary patterns of the EFNs: (1) change of habitat from forests to savannas (extrinsic factors); and (2) emergence of new morphological characters such as other trichome types over the plant\'s surface (intrinsic factors). Both factors might have altered the ant-plant interactions and the evolution of nectaries. The occupation of the savannas was associated with a decrease in the number of nectaries (likely due to counter-selection of these structures), while the evolution of adhesive glandular trichomes presenting the same effect on the nectaries. Both results are discussed in the light of the biotic variation (ants and herbivores) encountered between habitats, as well as in the light of the trade-off among defensive characters. The fourth chapter tested the geographic mosaic theory of coevolution in 10 populations of the savanna species Anemopaegma álbum. No correlations were found among extrafloral nectaries (and nectar variables), the abundance of visiting ants, herbivory, and plant performance among populations. This pattern was mainly associated with the variation in the assembly of ants encountered in the various populations. Most populations of A. álbum were dominated by assemblages of Camponotus ants, except for one that was dominated by Crematogaster ants. However, 3 of 10 populations studied presented a high number of plants without ants, decreasing the chances of defense against herbivores by EFNs. The abundance of ants was negatively associated with herbivory, and positively associated with plant performance variables among populations. Out of the 10 populations sampled, five presented an abundance of EFNs that matched the functional traits of ants. Out of these five populations, only three presented positive leaf production and low herbivory. From these three populations, two presented high abundances of EFNs on the leaves and were dominated by Camponotus ants (i.e., bigger in size, but with a low recruiting capacity). The third population presented on average the smallest abundance of EFNs on the leaves; it also presented the highest number of ants per plant (and the highest frequency), which generally were Crematogaster ants (smaller in size but with greater recruiting capacity). Because these ants are smaller in size, they used nearly all isolated EFNs encountered over the plant\'s surface. Neither the abundance of EFNs, the abundance of ants, and the assemblage of herbivores were structured spatially, corroborating the geographic mosaic hypothesis for the ant-plant-herbivore interactions in A. album. In this context, the three populations with \"matched\" ant-plant interactions were considered \'hot-spots\' of interactions, in which the populations reached the highest values of plant performance, while the others were considered \'cold-spots\'. The majority of \'cold spot\' populations were explained by a lack of sufficient ants to protect the plants effectively