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Made available in DSpace on 2016-04-25T16:44:52Z (GMT). No. of bitstreams: 1 Alexandre Marinho Teixeira.pdf: 841323 bytes, checksum: d2a82c49aabf948f5bfcf399070cd41b (MD5) Previous issue date: 2006-12-12 / The general objective of this work is to analyze the industrial sector of the Mate crop of the Southwest region of Paraná State. Analysis was performed through the value of the local productive arrangements and clusters, identifying their characteristics and specificity, links of partnerships and application of the concepts in the researched area. The mentioned area concentrates approximately 30% of the whole Mate crop industry of the State, corroborating with the concept of local productive arrangements. The research was accomplished among small, medium and big companies based in the region and members of FIEP (Federation of the Industries and Companies of the State of Paraná). The scale of Likert was used as the tool of measurement in order to identify the intensity degree of the actions used by the industries, such as the relationship level among the agents of a possible productive arrangement place. This work mentions, respectively, the history, the main approaches, interpretative chain and typologies of local productive arrangements and of clusters. As for the Mate crop industrial sector it portrays the extrativism in Brazil, the history, its uses, characteristics and improvement process, commercial movements and of the mate production. It makes an analysis, considerations and suggestions to contribute to the economical growth and development of the industrial Mate sector of the Southwest region of Paraná / A dissertação em questão tem como objetivo geral analisar o setor industrial ervateiro da região sudoeste do Estado do Paraná, por meio do valor dos arranjos produtivos locais e clusters, identificando suas características e especificidade, elos de parcerias e aplicação dos conceitos na região pesquisada. A região mencionada concentra aproximadamente 30% de toda a indústria ervateira do Estado do Paraná, corroborando com o conceito de arranjos produtivos locais. A pesquisa foi realizada entre empresas de pequeno, médio e grande porte sediadas na região e filiadas à FIEP (Federação das Indústrias e Empresas do Estado do Paraná). Utilizou-se como instrumento de mensuração a escala de Likert, a fim de identificar o grau de intensidade das ações empregadas pelas indústrias, como o nível de relacionamento entre os agentes de um possível arranjo produtivo local. O trabalho menciona os antecedentes históricos, as principais abordagens, correntes interpretativas e tipologias de arranjos produtivos locais e de clusters, respectivamente. Quanto ao setor industrial ervateiro retrata o extrativismo no Brasil, o histórico, usos, características e processo de beneficiamento, movimentos comerciais e de produção da erva mate. Realiza uma análise, considerações e sugestões que contribuem para o crescimento e desenvolvimento econômico do setor industrial ervateiro da região Sudoeste do Estado do Paraná

Among the different sensory modalities that play a role in sexual selection, acoustic communication plays an important one. Acoustic communication has been known to be used for male-male competition (territory maintenance, male aggression during mating),for advertisement to the opposite sex (mating status, body condition, genetic quality, nutritional status) and used by females to sample and choose conspecific preferred males. The use of acoustic communication for sexual display and information exchange has been extensively studied in multiple taxa, including insects, anurans, birds and mammals. Among insects, crickets have proven to be good model systems to study sexual selection based on acoustic communication as most species have an elaborate acoustic communication system, male advertisements, diverse types of mating incentives for females (such as glandular feeding) and a female dominated mating system. Generally, in crickets males produce species-specific calls which are used by females to localize conspecific males. Besides, calls show high levels of intraspecific variation and are energetically costly to produce. Moreover, as in crickets predominantly the females show phonotaxis towards male calls, calls also can play a role in mate sampling and choice by acting as indicators of preferred male quality. Despite being studied for many decades there are certain gaps in the studies examining mate choice in crickets. Some of them are, lack of understanding of the variation of male calling traits in nature and its role in signal evolution, lack of understanding regarding the ecological context of mate sampling and the evolution of alternative mating strategies. Hence, the tree cricket Oecanthus henryi was chosen as a study system to address these gaps in the understanding of female choice based on acoustic signals. In the tree cricket Oecanthus henryi, males call and females use calls to localize conspecific males and hence potentially females can choose males based on acoustic cues. To understand the evolution of female preference for male acoustic cues it is important to understand the variation in the calling songs in the field and identify repeatable call features that are reliable indicators of preferred male traits (morphological, developmental or genetic). I measured repeatability of male call traits in the field to understand their variation, reliability and consistency. Carrier frequency was the only call trait that was highly repeatable and hence was reliable and consistent. Following this I examined whether any of these call traits were indicators of male morphological traits (such as male size and fluctuating asymmetry) which are known to be preferred by females. It was found that carrier frequency was negatively correlated with body size; hence carrier frequency was both reliable and indicated male size. I also found that females preferred larger males during mating, as revealed by the longer mating durations and longer spermatophore retention time. Interestingly, though this study indicated that females could in principle use lower call carrier frequency to localize preferred larger males, simultaneous choice experiments done in the laboratory revealed that the females do not use this cue. These contrasting results may be because females are incapable of discriminating small differences in frequency or because they use non-acoustic cues for mate choice. However, whichever cues the females use to discriminate between males in the laboratory conditions, often these preferences are not realized in the field. The main reason behind this is that searching for preferred mates in the field can be costly and this might force females to choose sub-optimal males. Theoretical models predict that male movement and spacing in the field should influence female sampling tactics and in turn, females should drive the evolution of male movement and spacing to sample them optimally. Moreover, simultaneous sampling of males using the best-of-n or comparative Bayes strategy should yield maximum mating benefits to females. Many of the theoretical mate sampling strategies involves recall of the quality and location of individual males, which in turn requires male positions to be stable within a night. Calling males of O. henryi showed high site fidelity within a night, potentially enabling female sampling strategies that require recall. To examine the possibility of simultaneous acoustic sampling of males, I estimated male acoustic active spaces using information on male spacing, call transmission and female phonotactic threshold. Males were found to be spaced far apart and active space overlap was rare. I then examined female sampling scenarios by studying female spacing relative to male acoustic active spaces. Only 15% of sampled females could hear multiple males, suggesting that simultaneous mate sampling is rare in the field. Moreover, the relatively large distances between calling males suggest high search costs, which may favor threshold strategies that do not require memory. Using the insights gathered from these two studies I examined a unique calling behaviour from leaf holes, baffling, observed in this species. Baffling behaviour has been found in multiple species of the genus Oecanthus where the males call from selfmade holes in leaves rather than calling from leaf edges (their natural calling surface) thus increasing their loudness many fold. I started by examining the natural history of baffling and found that baffling is an extremely rare behaviour in the field. However field observations and laboratory experiments revealed that many males can baffle and hence it is not an obligatory behaviour shown only by a few males. It was hypothesized that one reason for the rarity of baffling could be resource limitation. It was found that baffling males prefer larger leaves possibly due to higher SPL gains achieved by baffling on the larger leaves, which is a limited resource in the field. However this alone was insufficient to explain extreme rarity of bafflers in the field. Hence I examined which males were using this behaviour in the field. Using field observations and laboratory experiments it was found that less preferred males (smaller and quieter) baffled more which provided them with higher calling SPL and greater sound-field volume and thus a higher number of potential mates. Moreover, baffling also increased the mating duration for the less preferred males thus providing more time to these males for sperm transfer. The females could not differentiate between an inherently loud caller and a caller whose SPL was increased artificially (as if it was baffling). Hence I concluded that baffling is probably a cheater strategy used by the less preferred males to fool the females into approaching them and mating for longer durations. To my knowledge, this is the first study that has estimated male call variation in the field to understand its role in female choice in tree crickets. Moreover this is also the first study to examine the ecological context of mate choice in tree crickets. This is also the first study to examine the advantages of baffling behaviour and its potential evolutionary implications. the different sensory modalities that play a role in sexual selection, acoustic communication plays an important one. Acoustic communication has been known to be used for male-male competition (territory maintenance, male aggression during mating),for advertisement to the opposite sex (mating status, body condition, genetic quality, nutritional status) and used by females to sample and choose conspecific preferred males. The use of acoustic communication for sexual display and information exchange has been extensively studied in multiple taxa, including insects, anurans, birds and mammals. Among insects, crickets have proven to be good model systems to study sexual selection based on acoustic communication as most species have an elaborate acoustic communication system, male advertisements, diverse types of mating incentives for females (such as glandular feeding) and a female dominated mating system. Generally, in crickets males produce species-specific calls which are used by females to localize conspecific males. Besides, calls show high levels of intraspecific variation and are energetically costly to produce. Moreover, as in crickets predominantly the females show phonotaxis towards male calls, calls also can play a role in mate sampling and choice by acting as indicators of preferred male quality. Despite being studied for many decades there are certain gaps in the studies examining mate choice in crickets. Some of them are, lack of understanding of the variation of male calling traits in nature and its role in signal evolution, lack of understanding regarding the ecological context of mate sampling and the evolution of alternative mating strategies. Hence, the tree cricket Oecanthus henryi was chosen as a study system to address these gaps in the understanding of female choice based on acoustic signals. In the tree cricket Oecanthus henryi, males call and females use calls to localize conspecific males and hence potentially females can choose males based on acoustic cues. To understand the evolution of female preference for male acoustic cues it is important to understand the variation in the calling songs in the field and identify repeatable call features that are reliable indicators of preferred male traits (morphological, developmental or genetic). I measured repeatability of male call traits in the field to understand their variation, reliability and consistency. Carrier frequency was the only call trait that was highly repeatable and hence was reliable and consistent. Following this I examined whether any of these call traits were indicators of male morphological traits (such as male size and fluctuating asymmetry) which are known to be preferred by females. It was found that carrier frequency was negatively correlated with body size; hence carrier frequency was both reliable and indicated male size. I also found that females preferred larger males during mating, as revealed by the longer mating durations and longer spermatophore retention time. Interestingly, though this study indicated that females could in principle use lower call carrier frequency to localize preferred larger males, simultaneous choice experiments done in the laboratory revealed that the females do not use this cue. These contrasting results may be because females are incapable of discriminating small differences in frequency or because they use non-acoustic cues for mate choice. However, whichever cues the females use to discriminate between males in the laboratory conditions, often these preferences are not realized in the field. The main reason behind this is that searching for preferred mates in the field can be costly and this might force females to choose sub-optimal males. Theoretical models predict that male movement and spacing in the field should influence female sampling tactics and in turn, females should drive the evolution of male movement and spacing to sample them optimally. Moreover, simultaneous sampling of males using the best-of-n or comparative Bayes strategy should yield maximum mating benefits to females. Many of the theoretical mate sampling strategies involves recall of the quality and location of individual males, which in turn requires male positions to be stable within a night. Calling males of O. henryi showed high site fidelity within a night, potentially enabling female sampling strategies that require recall. To examine the possibility of simultaneous acoustic sampling of males, I estimated male acoustic active spaces using information on male spacing, call transmission and female phonotactic threshold. Males were found to be spaced far apart and active space overlap was rare. I then examined female sampling scenarios by studying female spacing relative to male acoustic active spaces. Only 15% of sampled females could hear multiple males, suggesting that simultaneous mate sampling is rare in the field. Moreover, the relatively large distances between calling males suggest high search costs, which may favor threshold strategies that do not require memory. Using the insights gathered from these two studies I examined a unique calling behaviour from leaf holes, baffling, observed in this species. Baffling behaviour has been found in multiple species of the genus Oecanthus where the males call from self- made holes in leaves rather than calling from leaf edges (their natural calling surface) thus increasing their loudness many fold. I started by examining the natural history of baffling and found that baffling is an extremely rare behaviour in the field. However field observations and laboratory experiments revealed that many males can baffle and hence it is not an obligatory behaviour shown only by a few males. It was hypothesized that one reason for the rarity of baffling could be resource limitation. It was found that baffling males prefer larger leaves possibly due to higher SPL gains achieved by baffling on the larger leaves, which is a limited resource in the field. However this alone was insufficient to explain extreme rarity of bafflers in the field. Hence I examined which males were using this behaviour in the field. Using field observations and laboratory experiments it was found that less preferred males (smaller and quieter) baffled more which provided them with higher calling SPL and greater sound-field volume and thus a higher number of potential mates. Moreover, baffling also increased the mating duration for the less preferred males thus providing more time to these males for sperm transfer. The females could not differentiate between an inherently loud caller and a caller whose SPL was increased artificially (as if it was baffling). Hence I concluded that baffling is probably a cheater strategy used by the less preferred males to fool the females into approaching them and mating for longer durations. To my knowledge, this is the first study that has estimated male call variation in the field to understand its role in female choice in tree crickets. Moreover this is also the first study to examine the ecological context of mate choice in tree crickets. This is also the first study to examine the advantages of baffling behaviour and its potential evolutionary implications.

Cooperative breeding is a phenomenon whereby breeding and nonbreeding individuals collectively provision young. Nonbreeding group members ("helpers'') may gain indirect and/or direct fitness benefits by breeding in a group, but there has been conflicting evidence regarding the benefits to breeders. In fact, the presence of helpers may sometimes be detrimental to aspects of breeder fitness. For example, in some species of the chiefly Australian genus Malurus, breeding males with helpers have lower within-pair paternity than do males without helpers. Additionally, indirect benefits to breeding males are often limited by low relatedness to their helpers due to high extrapair paternity rates, and helpers often appear to have minimal impact on breeder reproductive success. However, the presence of helpers may allow breeding males to shift their behaviors from guarding and provisioning young to alternative behaviors that affect other components of fitness, such as extraterritory forays (which might increase extrapair mating success) and self-maintenance (which might increase survival). We investigated these possibilities in the facultatively cooperative Red-backed Fairywren (Malurus melanocephalus). Males with helpers spent significantly less time engaging in guarding behaviors and provisioning of young than did those without helpers, but there was no difference in the frequency of extrapair forays nor the number of young sired by males with vs. without helpers. Additionally, the decreased investment in nesting behaviors did not result in consistently higher survival, but may have increased survival in some years. Overall, the results of this study did not suggest any strong direct fitness benefits to breeding males, which may indicate that the costs of retaining helpers are negligible relative to the indirect benefits of helping a potentially related male.

Contrary to traditional belief, the decisions that females make before, during and after mating shape its outcome and ultimately fitness. The aim of this thesis was to examine how females modify and adjust their mating decisions in line with social and environmental variability and how these directly and indirectly affect mating benefits. To address this aim I have formulated four main questions that correspond to chapters 2 to 6 in this thesis. Firstly I asked whether there was evidence for female choice being driven by mating benefits. More precisely, in chapter 2, I reviewed the literature in search of evidence for direct and indirect benefits in female choice among freshwater fish species. Direct mating benefits were defined as an increase in female’s reproductive success (number of offspring). Conversely, increases in offspring reproductive success were considered to be indirect benefits. The results showed that despite the multiple suggestions and the great amount of information available, to date there is still no evidence for both direct (increase of F1) or indirect (increase in F2) mating benefits, nor their influence in female mating decisions if freshwater fishes species. Furthermore, although polyandry occurred in more than 60% of the species reviewed, I was unable to confirm that polyandry was maintained because of indirect benefits. These findings justified the need to experimentally investigate the drivers of female mating decisions in freshwater fish species and lead to the questions addressed on chapters 5 and 6. For the experimental chapters 3-6 I used the Trinidadian guppy, Poecilia reticulata, as a model species. Their unique and remarkable ecological and biological characteristics coupled with easy maintenance in laboratory conditions make guppies an ideal species for behavioural studies. But in particular, the fact that guppies live in a promiscuous mating system where females despite being constantly harassed by males may be able to control paternity, makes guppies the ideal species to address my aims. The second question was how much were females in control of their mating decisions, and how social environment could limit these decisions. In particular, in chapter 3 I looked at how females adapt and adjust their mating decisions in line with extreme differences in population sex ratio. In chapter 4, I examined to what extent male sexual harassment affects female reproductive behaviours. Results from these two chapters indicated that female guppies have a remarkable reproductive plasticity that enable them to control their mating decisions. When faced with extreme differences in population sex ratio, female mating decisions were made in an optimal way that maximized the ratio of female reproductive benefits per investment. This translated into producing bigger offspring when in a strong female biased environment, than when in a male biased environment. Further, chapter 4 illustrated that female guppies can, despite high levels of male sexual harassment, be in control of their mating decisions. These two chapters demonstrated and have reinforced previous findings of the remarkable reproductive adaptation of female guppies to differences in the social environment. The third question I addressed was: do multiply mated females have greater direct or indirect benefits than single mated females? To answer this question I followed for the first time reproductive success of females over two generations. I measured fitness directly (number of F1 and F2) and took as well as multiple indirect measures of fitness components for two generations in search for evidence of direct and indirect mating benefits in explaining the maintenance of female multiple mating. The results of chapter 5 revealed that female guppies do not have a higher number of F1 and F2 from polyandry or either from mating with males possessing allegedly good quality traits. I, therefore, stressed the idea that potential differences in sexual selection pressure between laboratory and wild populations may influence the expression and intensity of mating benefits between thus explaining the difficulty of finding mating benefits. In my fourth and last question, I used a novel statistical approach based on the analysis of the dispersion in phenotypes, to look for potential alternative explanations for the prevalence of polyandry. The results of this analysis show offspring from multiple mated mothers were phenotypically more diverse than offspring from single mated mothers. Given the direct relationship between phenotypic diversity and potential fitness gains in stochastic systems, female guppies are likely to get greater benefits from mating with males with different phenotypes than with males with a particular sexual trait. This result provides an alternative explanation for the maintenance of polyandry in resource free systems. Overall the results of this thesis reinforce previous suggestions that female guppies are active participants in the mating process, and not necessarily limited to post-copulatory mechanisms of selection of sperm. It also showed the remarkable ability of females to adjust their reproductive investment in line with changes in the social conditions. Interestingly, my results contradict the commonly accepted assertion that females’ mating preference converges towards unique male sexual traits. This result stresses the need to look at alternative explanations to justify female mating decisions.

Learning and experience shape mate preferences in many species. My thesis investigates the role of experience on mating behavior of male and female sailfin mollies (Poecilia latipinna). In the first chapter I explore whether adult experience influences male sailfin molly mate preference for their sexual parasite, the Amazon molly (Poecilia formosa), and whether experience could account for reproductive character displacement (RCD) of male mate preference in this species. Sailfin males from sympatric populations show a stronger preference for conspecific females over Amazon mollies than do males from allopatric populations. I exposed males from sympatric and allopatric populations to either a sailfin female or an Amazon prior to a mating trial with an Amazon. For the allopatric population, males with recent experience with an Amazon directed fewer mating behaviors towards an Amazon during mating trials than did males with recent experience with a sailfin. Males from the sympatric population, however, performed the same amount of mating behaviors towards an Amazon regardless of experience. Thus adult experience influences mating preferences and suggests that experience may play a role in RCD in this species. In the second chapter I investigate whether a learned sensory bias could influence female mate preferences. Sensory biases that influence mate preferences can arise through selection on the sensory system in foraging and predator detection domains. I tested whether a learned preference originating outside of the mating domain, specifically a color-based food preference, can be transferred to a color-based preference for a male trait. I trained female sailfin mollies to associate either green or blue with food and then tested their preference for animated male sailfins featuring either a blue or green spot. I found that females did not prefer the male with the same color spot to which they had been conditioned. I discuss the problem of learned preference transfer and suggest directions for future research into the role of learning in sensory bias. / text

Despite recent increasing interest in the existence of animal personality, i.e. intra-individual consistency and inter-individual variation in the level of a behavioural trait, the evolutionary (and ecological) consequences of these consistent behavioural differences remain poorly understood. Some recent studies have revealed that variation in animal personalities might be linked to competitive interactions, resulting from natural selection. However, since personalities might similarly affect mate acquisition and reproductive success, it seems crucial to also explore their evolution under the framework of sexual selection theory. In this thesis I investigate the influence of personality on mate choice, reproductive success, female-male and male-male interactions, using zebra finches, Taeniopygia guttata, as a model species. After (I) I review the few existing studies assessing the link between sexual selection and personality, I propose a framework on the relationship between sexual selection and personality. Subsequently, (II) I show with different experimental approaches for the first time that (a) females choose males on the basis of their behaviour per se (male behaviour was experimentally disentangled from any appearance effects) whilst considering their own personality in their choice: less exploratory females did not distinguish between exploratory and non-exploratory appearing males, whereas moderately and highly exploratory females preferred similar males. (b) These preferences have an adaptive value to the exploratory females: exploratory females which had a behaviourally similar partner raised chicks in best condition but chicks in worst condition if they had a dissimilar partner. Low exploration females always raised chicks in intermediate condition, which may explain why they did not choose males on the basis of their exploratory behaviour. (c) I provide evidence that the combination of personalities in a pair, not only in terms of the level of the behaviour but also in terms of the behavioural consistency, influence reproductive success. However, this was only true for foster but not genetic parents, suggesting that behavioural rather than genetic compatibility (for the measured personality traits) is important for reproduction. (d) Moreover, some male behavioural characteristics appear to be a signal of male quality: highly exploratory foster males raised chicks (both males and females) in best condition, which themselves raised foster chicks of increased conditions. (e) Furthermore, a number of sex differences in personality traits (both level and consistency) are identified, including different responses to social interactions. (f) Additionally, I show how females with different exploratory tendencies differ in their movement patterns during mate choice. (g) Finally, I demonstrate that in competitive male-male situations, a measurement of condition, the fat score, and aggressive behaviour are positively correlated. These findings are set into sexual selection context (but other evolutionary processes are also considered) and both their ecological and evolutionary consequences are discussed. I outline how these results make a valuable contribution to the research field and discuss their potential to indicate new directions for future studies.

Over the last century, sexual selection has grown from a controversial theory into a vast field of theoretical and empirical research. Although Darwin outlined two major mechanisms within his theory, male-male competition and female mate choice, the latter has promoted a wealth of research by virtue of its complexity. Despite decades of research into how female preferences and sexually selected traits have evolved, there is still little consensus as to why females prefer the males they do. Preferences are thought to evolve from either direct selection on the preference, as females themselves benefit directly from mating with a preferred male, or through indirect selection on the preference via offspring fitness. In all cases however, female preferences should compensate for the costs of discriminating between potential mates, if they are to remain overall beneficial. The fitness benefits of mating with preferred males were investigated here using the fruitfly Drosophila simulans, employing a range of behavioural, phenotypic and quantitative genetic approaches. The findings presented here indicate that female Drosophila simulans do not gain directly from mating with a preferred male. Multiple mating can increase fecundity, although costs from male harassment can reduce the net benefit. They also indicate that females may benefit indirectly from mating with attractive males as attractiveness is heritable and sons of preferred males are themselves preferred. There is also evidence that attractive males are successful in both the pre- and post-copulatory sense, as preferred males are better sperm competitors than less-preferred males. However, although there appear to be benefits from preferred males via their sons, there appear to be no benefits from males via their daughters’ fitness. These findings collectively indicate that female preferences in Drosophila simulans are driven by indirectly selected benefits (via Fisherian sons), and that females benefit directly from mating multiply.