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<p>Alternative reproductive tactics (ARTs) have been described across taxa numerous times, and are especially common in fishes. The advent of molecular techniques has enhanced our understanding of such mating systems, yet these techniques have only been applied to relatively few species. Furthermore, ecological variation has long since been recognized as an important factor influencing mating patterns and sexual selection, yet it is often ignored. In my thesis, I conducted field studies to investigate the reproductive ecology of a species with well characterized ARTs, the plainfin midshipman (<em>Porichthys notatus</em>), and to explore how these mating patterns varied across time and space. This species is characterized by two male tactics: guarders and sneakers. I show that both tactics are successful at gaining fertilization. Indeed, I found evidence for several behavioural adaptations by the guarder male tactic to gain fitness, including guarding, nest takeovers, and plastic cuckoldry behaviours. I also documented the lowest paternity for guarder males for a species with obligate male parental care. Paternity increased over the course of the breeding season, which was likely a result of a higher occurrence of nest takeovers early in the season. In a cross-population analysis of plainfin midshipman, I found significant differences in nest availability and density between two genetically distinct populations, but this did not translate into significant differences in mating patterns. Taken together, my thesis helps elucidate the reproductive ecology of a classic species with ARTs, and has important implications for our understanding of ecological influences on mating patterns and sexual selection.</p> / Doctor of Philosophy (PhD)

Ainda que existam diversos registros de formas extremas de dimorfismo sexual, a função do alongado quarto par de pernas nos machos de muitas espécies de opiliões permanece desconhecida. Neste estudo, testamos quatro previsões sobre a hipótese de que a seleção sexual está direcionando o alongamento do quarto par de pernas nos machos de Longiperna zonata e Promitobates ornatus: (1) o coeficiente alométrico do comprimento do fêmur IV em machos será maior do que o das fêmeas; (2) machos com pernas IV mais compridas vão residir próximos a grupos de fêmeas; (3) machos envolvidos em interações agressivas terão fêmures mais compridos do que o tamanho médio da população; (4) a diferença de tamanho de perna entre os oponentes será menor do que a diferença esperada entre pares de machos escolhidos ao acaso na população. Como o previsto, o coeficiente alométrico do fêmur IV foi significativamente maior em machos do que em fêmeas nas duas espécies. O comprimento do fêmur IV está positivamente relacionado à chance de um macho ser encontrado próximo a um grupo de fêmeas. Finalmente, a média do comprimento do fêmur IV dos machos envolvidos em brigas foi significativamente maior que a de machos selecionados ao acaso da população. Adicionalmente, a média da diferença de tamanho entre pares de machos que brigaram foi significativamente menor que a média da diferença de tamanho de perna entre pares de machos selecionados ao acaso na população. Assim, a seleção sexual na forma de competição entre machos parece estar direcionando o dimorfismo sexual extremo no comprimento do fêmur nessas duas espécies de opiliões. / Although there are many reports of extreme forms of sexual dimorphism in harvestmen, the function of the elongated fourth pair of legs in males of many species remains unknown. We tested four predictions to address the hypothesis that sexual selection is driving the enlargement of the fourth pair of legs in males of Longiperna zonata and Promitobates ornatus: (1) the allometric coefficient of femur IV length in males will be higher than in females; (2) males with longer legs IV will reside closer to groups of females; (3) males involved in aggressive interactions will have longer femur IV than the population average size; (4) the size difference between contenders will be smaller than the difference expected between randomly chosen pairs of males in the population. As predicted, the allometric coefficient of femur IV length was significantly higher in males than in females of both species. Femur IV length was positively related to the chance of a male being found close to a group of females. Finally, the mean femur IV length of the males involved in fights was significantly larger than the mean of random samples of males from the population, and the average difference between male contenders was smaller than the average random expected difference of the population. Therefore, sexual selection in the form of male-male competition seems to be driving the extreme sexual dimorphism in femur length in these species.

Darwinian sexual selection predicts that males with the most extravagant secondary sexual traits suffer elevated mortality. Although correlative evidence has generally not borne this idea out, recent research, including a field cricket study, showed that investment in sexually selected traits is costly to survival. I investigated male survival, ornamentation (song) and mating success in a North American grylline, Gryllus pennsylvanicus, to test the generality of previous work and highlight the importance of ecology differences to resource allocation. As the calling songs of older male G. pennsylvanicus are highly attractive to females, in Chapter 2 I tested whether male age correlated with calling song and found a weak but statistically significant correlation, thus leaving open the possibility that choosy females use an age-based indicator mechanism. In Chapter 3, I tested the condition dependence of male survival and calling effort. In contrast to previous work, I found that high condition males both called more and lived longer than low condition males, although there was no trade-off between survival and calling effort. The substantial condition dependence of calling effort suggests that calling effort is under strong directional selection. In Chapter 4 I tested whether female mating preferences resulted in strong selection on male calling effort. I also tested for the condition dependence of female mating preferences. I found that female choosiness was condition-dependent, but the rank of preferred male songs (preference function) was not. Both low and high condition females preferred high calling effort over low calling effort song. In Chapter 5 I tested for evidence of nonlinear selection on male survival that might explain the nonlinear pattern of male investment in survival seen in Chapter 3 (i.e. male survival leveled-off with increasing condition). I found that socially experienced females, but not virgin and naive females, exerted linear selection on male age. I discuss these and the other results of my thesis in the context of previous work on field crickets and condition-dependent ornamentation. Finally, Appendix A reports results that confirm ancient Chinese cultural knowledge that large headed male crickets are more successful in male-male combat.

Darwinian sexual selection predicts that males with the most extravagant secondary sexual traits suffer elevated mortality. Although correlative evidence has generally not borne this idea out, recent research, including a field cricket study, showed that investment in sexually selected traits is costly to survival. I investigated male survival, ornamentation (song) and mating success in a North American grylline, Gryllus pennsylvanicus, to test the generality of previous work and highlight the importance of ecology differences to resource allocation. As the calling songs of older male G. pennsylvanicus are highly attractive to females, in Chapter 2 I tested whether male age correlated with calling song and found a weak but statistically significant correlation, thus leaving open the possibility that choosy females use an age-based indicator mechanism. In Chapter 3, I tested the condition dependence of male survival and calling effort. In contrast to previous work, I found that high condition males both called more and lived longer than low condition males, although there was no trade-off between survival and calling effort. The substantial condition dependence of calling effort suggests that calling effort is under strong directional selection. In Chapter 4 I tested whether female mating preferences resulted in strong selection on male calling effort. I also tested for the condition dependence of female mating preferences. I found that female choosiness was condition-dependent, but the rank of preferred male songs (preference function) was not. Both low and high condition females preferred high calling effort over low calling effort song. In Chapter 5 I tested for evidence of nonlinear selection on male survival that might explain the nonlinear pattern of male investment in survival seen in Chapter 3 (i.e. male survival leveled-off with increasing condition). I found that socially experienced females, but not virgin and naive females, exerted linear selection on male age. I discuss these and the other results of my thesis in the context of previous work on field crickets and condition-dependent ornamentation. Finally, Appendix A reports results that confirm ancient Chinese cultural knowledge that large headed male crickets are more successful in male-male combat.

In this study I investigate for two consecutive breeding seasons male calling patterns in natural population of European treefrog (Hyla arborea), a hylid frog with prolonged breeding season and a lek mating system. I tested the hypothesis that males modify their vocal behavior in different ways when faced with stimuli of competitors with various body size. Further here, the influence of water temperature and size of advertising males on their call properties were concerned.

Ainda que existam diversos registros de formas extremas de dimorfismo sexual, a função do alongado quarto par de pernas nos machos de muitas espécies de opiliões permanece desconhecida. Neste estudo, testamos quatro previsões sobre a hipótese de que a seleção sexual está direcionando o alongamento do quarto par de pernas nos machos de Longiperna zonata e Promitobates ornatus: (1) o coeficiente alométrico do comprimento do fêmur IV em machos será maior do que o das fêmeas; (2) machos com pernas IV mais compridas vão residir próximos a grupos de fêmeas; (3) machos envolvidos em interações agressivas terão fêmures mais compridos do que o tamanho médio da população; (4) a diferença de tamanho de perna entre os oponentes será menor do que a diferença esperada entre pares de machos escolhidos ao acaso na população. Como o previsto, o coeficiente alométrico do fêmur IV foi significativamente maior em machos do que em fêmeas nas duas espécies. O comprimento do fêmur IV está positivamente relacionado à chance de um macho ser encontrado próximo a um grupo de fêmeas. Finalmente, a média do comprimento do fêmur IV dos machos envolvidos em brigas foi significativamente maior que a de machos selecionados ao acaso da população. Adicionalmente, a média da diferença de tamanho entre pares de machos que brigaram foi significativamente menor que a média da diferença de tamanho de perna entre pares de machos selecionados ao acaso na população. Assim, a seleção sexual na forma de competição entre machos parece estar direcionando o dimorfismo sexual extremo no comprimento do fêmur nessas duas espécies de opiliões. / Although there are many reports of extreme forms of sexual dimorphism in harvestmen, the function of the elongated fourth pair of legs in males of many species remains unknown. We tested four predictions to address the hypothesis that sexual selection is driving the enlargement of the fourth pair of legs in males of Longiperna zonata and Promitobates ornatus: (1) the allometric coefficient of femur IV length in males will be higher than in females; (2) males with longer legs IV will reside closer to groups of females; (3) males involved in aggressive interactions will have longer femur IV than the population average size; (4) the size difference between contenders will be smaller than the difference expected between randomly chosen pairs of males in the population. As predicted, the allometric coefficient of femur IV length was significantly higher in males than in females of both species. Femur IV length was positively related to the chance of a male being found close to a group of females. Finally, the mean femur IV length of the males involved in fights was significantly larger than the mean of random samples of males from the population, and the average difference between male contenders was smaller than the average random expected difference of the population. Therefore, sexual selection in the form of male-male competition seems to be driving the extreme sexual dimorphism in femur length in these species.

A critical mechanism underpinning current biological diversity is the extent to which one species mates with, or avoids mating with, another. However, little is known about the factors that mediate hybridisation, especially during the initial and rarely observed stages of secondary contact when interspecific interactions have not responded to selection. In particular, whilst hybridisation is ultimately a behavioural phenomenon, the role of behaviour in mediating hybridisation and how it is influenced by environmental and circumstantial factors is rarely investigated. Recently introduced species provide us with unequalled opportunities to study these factors. In this thesis I examine the role of behavioural mechanisms, in particular male-male competition and mate choice, in mediating mating patterns between two genetically and phenotypically distinct lineages of European wall lizard (Podarcis muralis) that have come into recent secondary contact through human introductions. In Chapter Two, I investigated how sexual selection during allopatry is responsible for creating stark differences in phenotypic traits such as body size and weapon performance evident in the two lineages today, ultimately explaining the strong biases in dominance during territorial disputes between males. However, I also show that even given this asymmetry in male competitive ability, the extent to which it extrapolates into greater access to females in naturalistic, outdoor enclosures depends strongly on the spatial clustering of basking sites, a critically important resource for many ectotherms. In contrast to initial predictions suggested by asymmetries in male competition outlined in the previous chapter, in Chapter Three I show that both paternity and courtship behaviour was strongly assortative in the outdoor enclosures. Further investigation through staged experiments on olfactory mate choice, mating trials and analyses on specific behavioural data obtained in an enclosure experiment, I show that lineage based dominance actually contributes to assortative mating patterns in conjunction with weak conspecific male choice. In contrast, female choice seems to play no role in mediating the mating patterns observed between the two lineages. In Chapter Four I had the rare opportunity to examine the morphological and behavioural factors that predict why animals should hybridise in the first place, using the data obtained in the enclosure experiment above. I found that hybridisation was particularly common between small individuals of the larger lineage and large individuals of the smaller lineage; a result that corroborates the mechanisms determining the assortative patterns uncovered in Chapter Three. Additionally, hybridisation rates were particularly high in less dominant individuals, which I suggest is due to subordinate males having reduced opportunities for courting conspecific females due to male-male competition, requiring them to become less ‘choosy’ and therefore more likely to mate with heterospecifics. Finally, secondary contact cannot occur without at least one lineage coming into a new environment, and yet relatively little attention is paid to how this environmental change can affect the signals involved in intraspecific communication and mate choice. In Chapter Five I show that a change in the amount of time male lizards spend thermoregulating (a likely consequence of arriving in a new environment) significantly changes the chemical composition of their scent marks. However, whilst female lizards were able to detect these effects, they did not seem to base their mating decisions on them. Nevertheless, this result raises interesting questions about the potential function and consequences of this plasticity, and highlights the importance of considering plasticity in chemical communication in heterogeneous environments. Overall, this thesis shows the critically important role of behaviour in mediating intra- and interspecific mating patterns during recent secondary contact. In particular, it highlights how the direction and extent of hybridisation and competition are influenced by the degree to which differing morphological and behavioural phenotypes interact over a heterogeneous environment, particularly during the initial stage of secondary contact when mate choice has not had the chance to respond to the selective pressures of hybridisation.

The availability of reproductive sites is a major factor shaping the behavior of males and females in species with resource-based mating systems. Using a meta-analytic approach, we tested five predictions directly or indirectly derived from the mating system theory. We expected that reproductive site limitation would lead to: (1) intense male-male competition for resource possession; (2) high variance in male reproductive success, generating high values of opportunity for sexual selection; (3) high intensity of selection on male traits related to resource possession; (4) high sperm competition risk; and (5) low quality of paternal care. We compiled information from observational and experimental studies that compared the reproductive behavior of individuals of the same species under low and high reproductive site limitation. We found that, when reproductive sites are limited, there is a slight increase in male-male competition, with higher rates of nest takeover and agonistic interactions, and a slight increase in the selection gradient on male traits, with successful males tending to be larger than unsuccessful males. Reproductive site limitation has no consistent effect on the opportunity for sexual selection and on the sperm competition risk. However, territorial males invest more in gonads and lose less paternity when reproductive sites are limited. There is also no clear effect of reproductive site limitation on the quality of paternal care, but few studies have addressed this subject. Taken together, our findings indicate that predictions on how reproductive site limitation affects several aspects of resource-based mating systems have weak empirical support. These predictions do not consider the plasticity in the mating tactics of males and females, which make them too simplistic. Moreover, since the original proposition of the mating system theory, our understanding of sperm competition increased a lot. We now know that accurate predictions on the intensity and direction of sexual selection should take into account both pre- and post-copulatory processes. Finally, the interplay between sexual selection and parental care is complex, and the original framework of mating system theory does not provide sufficient elements to derive clear and taxonomically broad predictions / A disponibilidade de sítios reprodutivos influencia tanto o comportamento de machos quanto de fêmeas em espécies cujo sistema de acasalamento depende da defesa de recursos. Usando uma abordagem meta-analítica, testamos cinco previsões direta ou indiretamente relacionadas à teoria de sistemas de acasalamento. Esperávamos que a limitação de sítios reprodutivos promoveria: (1) aumento na competição masculina pela posse de recursos; (2) aumento na variância do sucesso reprodutivo dos machos, gerando valores alto de oportunidade para a seleção sexual; (3) aumento da intensidade da seleção sobre características masculinas relacionadas à posse de recursos; (4) aumento no risco de competição espermática e (5) diminuição da qualidade do cuidado paternal. Compilamos informações de estudos observacionais e experimentais que compararam o comportamento reprodutivo de indivíduos da mesma espécie em situação de alta e baixa disponibilidade de sítios reprodutivos. Encontramos que, quando os sítios reprodutivos são escassos, há um ligeiro aumento na competição masculina, com maiores taxas de roubo de ninhos e interações agonísticas, e um ligeiro aumento no gradiente de seleção sobre características masculinas, com machos bem sucedidos tendendo a ser maiores do que machos mal sucedidos em monopolizar recursos. A disponibilidade de sítios reprodutivos não teve nenhum efeito consistente sobre a oportunidade para a seleção sexual e o risco de competição espermática. Entretanto, machos territoriais investiram mais em gônadas e perderam menos paternidade quando os sítios reprodutivos eram escassos. Não encontramos nenhum efeito da disponibilidade de sítios reprodutivos sobre a qualidade do cuidado paternal. Em conjunto, nossos resultados indicam que as previsões sobre como a disponibilidade de sítios reprodutivos influencia diversos aspectos de sistemas de acasalamento baseados na defesa de recursos têm fraco respaldo empírico. Tais previsões não consideram a plasticidade nas táticas de acasalamento de machos e fêmeas, tornando-as demasiadamente simplistas. Adicionalmente, desde a formulação original da teoria de sistemas de acasalamento, nossa compreensão sobre competição espermática aumentou. Sabemos atualmente que previsões acuradas sobre a intensidade e direção da seleção sexual devem levar em consideração processos pré- e pós-copulatórios. Finalmente, a interconexão entre seleção sexual e cuidado parental é complexa e a teoria original de sistemas de acasalamento não provê elementos suficientes para a construção de previsões claras e de amplo escopo taxonômico

Dans un contexte de sélection sexuelle, les systèmes de communication permettant l’attraction et la stimulation du partenaire sexuel et la compétition entre individus du même sexe sont indispensables. Ceci est particulièrement vrai chez les espèces à système d’appariement polygyne de type lek, où les mâles rassemblés dans l’espace sont en compétition pour l’accès aux femelles. Chez notre sujet d’étude, l’outarde houbara nord-africaine les males réalisent des parades incluant des démonstrations visuelles et des vocalisations appelées booms, sur des sites espacés les uns des autres par des distances importantes dans un système qualifié de lek explosé. Notre objectif était d’étudier les systèmes de codage-décodage des informations exprimées durant la parade des mâles chez cette espèce. Nous avons démontré l’existence d’un codage de l’identité individuelle porté par des paramètres visuels et des paramètres acoustiques des vocalisations. Des associations significatives entre des paramètres des booms et des caractéristiques physiques et comportementales des mâles susceptibles de refléter leur qualité, ont également été mises en évidence. Ainsi, les mâles qui produisent les booms les plus graves avaient les masses les plus importantes et paradaient le plus intensément. Lorsque des interactions agonistiques étaient simulées expérimentalement en diffusant des leurres acoustiques, les mâles avaient des réponses comportementales différentes en fonction de la fréquence des booms diffusés. Ainsi le niveau fréquentiel des booms des autres mâles est effectivement perçu et décodé par les individus en compétition. Par ailleurs, plusieurs paramètres des booms sont génétiquement déterminés et héritables, et pourraient porter une information sur l’apparentement entre individus.Certains paramètres sont également liés à la consanguinité des individus. Nous avons montré également que les booms sont particulièrement bien adaptés à la communication à grande distance. En effet, les booms sont de très basse fréquence, se propagent à des distances supérieures à 640 m, et les paramètres supportant le codage-décodage de l’information sont résistants à la propagation et produits de manière redondante. Enfin, les signaux acoustiques et des signaux visuels et semblent agir en synergie (multimodalité) ce qui pourraient améliorer l’efficacité de la communication à grande distance. Ainsi notre étude a démontré que des informations complexes sont transmises durant la parade des mâles au travers de signaux acoustiques et visuels au sein d’un réseau de communication à grande distance. / Résumé en anglais ;In a context of sexual selection, the communication between individuals is essential to optimize both intra-sexual (competition between male) and inter-sexual (mate choice) processes. This is particularly true in species with lek mating systems, where both processes reach their full expression. The North African Houbara bustard, Chlamydotis undulata undulata, is a lekking bird where courting males aggregate in a so called ‘exploded-lek’, i.e. where aggregation is detectable only when mapped over a large area. During the breeding season, males perform a conspicuous and spectacular courtship including visual and acoustic (booms) signals. Our aim was to decipher the coding-decoding systems of the courtship in this species. We found that males can be individually discriminated on the basis of visual and acoustic parameters. The booms produced during the courtship are also related to males’ characteristics susceptible to reflect their quality. Individuals who produced the lowest frequency booms were those with the greatest weights and performed the highest intensity courtships. Playback experiment shows that the frequency content of the booms was perceived and decoded by males during agonistic interaction, leading to significantly different behavioural responses according to the frequency of the signals played back. Furthermore, several acoustic parameters of the booms are significantly heritable through genetic transmission and these parameters are also related to the level of inbreeding. Finally, we showed that the booms and the acoustic parameters supporting information are particularly well adapted to the transmission of information at long range as required in a long distance communication network constituted by an exploded lek. First, booms are of very low-frequency and propagate up to 640 m. Second, coding-decoding of information relies on sequentially redundant and propagation-resistant features. Third, acoustic signals are combined with visual signalling (multimodality) which may improve the long distance transmission of the information. This study brought evidence that complex information is conveyed by visual and acoustic signals during the courtship of males within a long distance network.